Albuquergue Et Al Schizoptera Zygophleps

Insect Systematics & Evolution (2022) DOI 10.
1163/1876312X-bja10029
Taxonomic revision of the minute litter bug subgenus

Schizoptera (Zygophleps) McAtee & Malloch, 1925
(Heteroptera: Dipsocoromorpha: Schizopteridae)
Flavio Albuquerquea,b,*,**, Jose A.M. Fernandesa,*** and Christiane Weirauchb,****

aDepartment of Zoologia ‒ Invertebrados, Instituto de Ciências Biológicas, Universidade Federal
do Pará, Belém, Pará, 66075–110, Brazil
bDepartment of Entomology, University of California Riverside, Riverside, CA 92521, USA
*Corresponding author, e-mail: flavio.heteroptera@gmail.com
**ORCID: 0000-0001-5711-7259
***ORCID: 0000-0001-7450-5296
****ORCID: 0000-0003-4492-4515
Abstract
Schizoptera Fieber is a monophyletic and the largest genus of minute litter bugs (Hemiptera: Heteroptera:
Dipsocoromorpha). Its current concept is supported by a recent molecular phylogeny and comprises
nearly 90 described extant and two fossil species divided into four subgenera: Schizoptera (Cantharocoris),
Schizoptera (Odontorhagus), Schizoptera (Schizoptera) and Schizoptera (Zygophleps). We here taxonomi-
cally revise the subgenus Schizoptera (Zygophleps), which so far contains Schizoptera unica McAtee &
Malloch, 1925 from Guatemala and Schizoptera corallia Emsley, 1969, Schizoptera simla Emsley, 1969,
and Schizoptera ultima Emsley, 1969 from Trinidad. The subgenus is diagnosed by the unique triangular
shape of the posterior membranal cell. The following 36 species are here described: Schizoptera aculeata
sp.n., Schizoptera applanata sp.n., Schizoptera bella sp.n., Schizoptera biancae sp.n., Schizoptera cassiae
sp.n., Schizoptera confusa sp.n., Schizoptera costarriquenha sp.n., Schizoptera cuspida sp.n., Schizoptera
ecuadoriana sp.n., Schizoptera elegantissima sp.n., Schizoptera emsleyi sp.n., Schizoptera erosa sp.n.,
Schizoptera erwini sp.n., Schizoptera falcata sp.n., Schizoptera familia sp.n., Schizoptera furcata sp.n.,
Schizoptera fuscodorsata sp.n., Schizoptera hirsuta sp.n., Schizoptera inaequalis sp.n., Schizoptera inca sp.n.,
Schizoptera kayi sp.n., Schizoptera knyshovi sp.n., Schizoptera magnifica sp.n., Schizoptera mallochi sp.n.,
Schizoptera mcateei sp.n., Schizoptera orellanensis sp.n., Schizoptera paraensis sp.n., Schizoptera peruviana
sp.n., Schizoptera pichincha sp.n., Schizoptera pinduca sp.n., Schizoptera prima sp.n., Schizoptera priva
sp.n., Schizoptera ramosa sp.n., Schizoptera solitaria sp.n., Schizoptera speira sp.n., and Schizoptera stricta
sp.n. The majority of species herein described were collected in Brazil, Costa Rica, and Ecuador, expand-
ing the known distribution of the subgenus. We provide diagnoses, descriptions, photographs of habitus
and subgenital plates, line drawings of diagnostic male genitalic features, and a distribution map. A key to
species based on males is also presented.
© Koninklijke Brill NV, Leiden, 2022DOI 10.1163/1876312X-bja10029

F. Albuquerque et al. / Insect Systematics & Evolution (2022)
2 DOI 10.1163/1876312X-bja10029
Keywords
Hemiptera; Neotropical region; new species; Schizopterinae; taxonomy
ZooBank: http://zoobank.org/D1D2E0A6-84E6-4855-AF29-2F7F9F0BB20A
Introduction
The cosmopolitan true bug infraorder Dipsocoromorpha, usually referred as the min-
ute litter bugs, is one of the smallest and least well documented taxa of Heteroptera,
comprising about 430 species divided into six families (Schuh & Weirauch 2020).
Historically, this group was composed of the families Ceratocombidae, Dipsocoridae,
Hypsipterygidae, Schizopteridae, and Stemmocryptidae. Based on recent molecular and
morphological phylogenetic analyses, Trichotonanninae that was previously regarded as a
ceratocombid subfamily, are now treated at the family level (Knyshov et al. 2020).
Schizopteridae is the best known dipsocoromorphan family composed of approx-
imately 360 species divided into around 60 genera (Weirauch & Fernandes 2015;
Weirauch et al. 2018; Almeida et al. 2019). Specimens are tiny (~1–2 mm) and typ-
ically found in cryptic microhabitats such as leaf litter, low herbaceous vegetation,
under bark of fallen trees, and in the canopy (Emsley 1969; Weirauch & Fernandes
2015; Weirauch & Štys 2014). Species in several genera are attracted to light traps but
can also be abundant in bulk samples of other types of passive traps, including leaf
litter extraction and Malaise traps. Since specimens are very small and require special-
ized curation, most have remained uncurated, making them inaccessible for species
discovery and documentation.
During the past few years, the Weirauch laboratory has retrieved, mostly from bulk
samples, more than 10,000 schizopterid specimens from numerous institutions around
the world. More than 3,300 of these specimens belong to the genus Schizoptera Fieber,
1860 (Leon & Weirauch 2016a), the largest dipsocoromorphan genus worldwide, and
the dominant group of schizopterids in the New World.
The current concept of Schizoptera comprises nearly 90 described extant and two
fossil species divided into four subgenera: S. (Cantharocoris), S. (Odontorhagus), S.
(Schizoptera), and S. (Zygophleps) (Leon & Weirauch 2016a; Leon & Weirauch 2016b;
Leon & Weirauch 2017). Leon & Weirauch (2017) proposed the first phylogenetic
hypothesis for Schizoptera and related genera based on molecular data. The results
showed that the previous concepts of the genus used by McAtee & Malloch (1925) and
Emsley (1969) were polyphyletic. Therefore, Leon & Weirauch (2016a, 2017) raised
Kophaegis McAtee & Malloch and Orthorhagus McAtee & Malloch to genus level,
and synonymized S. (Lophopleurum) with S. (Cantharocoris), rendering Schizoptera
monophyletic. Moreover, Leon & Weirauch (2017) showed that the subgenera
S. (Cantharocoris), S. (Odontorhagus), and S. (Zygophleps) are monophyletic whereas
S. (Schizoptera) is polyphyletic and highlighted the importance of taxonomic revisions
across the genus.
The subgenus S. (Zygophleps) is currently composed of S. (Zygophleps) unica McAtee
& Malloch, 1925, S. (Zygophleps) simla Emsley, 1969, S. (Zygophleps) corallia Emsley,
DOI 10.1163/1876312X-bja10029 3
1969 and S. (Zygophleps) ultima Emsley, 1969. It is only known from Guatemala and
Trinidad. The subgenus is easily distinguished from other Schizoptera subgenera by the
unique triangular posterior membranal cell. More than 200 specimens of Schizoptera
(Zygophleps) from several countries including Brazil, Costa Rica, and Ecuador have
become available as part of the above-mentioned bulk sample sorting efforts, and addi-
tional specimens were collected in the field in the Brazilian Amazon by the first author.
Preliminary examination suggested that this material contained several undescribed
species; similar to other projects focused on Schizopteridae, it only became clear after
careful examination of specimens that this relatively small number of specimens repre-
sent a ten-fold increase of described species in this subgenus.
We here taxonomically revise the subgenus S. (Zygophleps), redescribe previously
known species and describe 36 species as new. Since species of this subgenus are
superficially very similar the diagnoses rely heavily on features of the male pregeni-
tal abdomen and male genitalia (e.g., subgenital plate, right conjunctival appendage).
Additionally, we provide a key to species based on males, digital images of habitus and
subgenital plates, and line drawings of genitalic diagnostic features. A distribution map
for species of the subgenus is also provided.
Material and nethods

Material
We examined 241 specimens, 51 of which were point-mounted and 190 preserved
in ethanol. The specimens utilized in this study were taken on loan from and will be
deposited in the following institutions:
AMNH American Museum of Natural History, New York, USA
FMNH The Field Museum of Natural History, Chicago, USA
FSCA Florida State Collection of Arthropods, Gainesville, USA
INBio Instituto Nacional de Biodiversidad, San Jose, Costa Rica
LACM Natural History Museum of Los Angeles County, Los Angeles, USA
MPEG Museu Paraense Emílio Goeldi, Belém, Brazil
MTEC Montana Entomology Collection, Bozeman, USA
TAMU Texas A&M University, College Station, USA
UCD University of California, Davis, USA
UCR University of California, Riverside, USA
UFPA Universidade Federal do Pará, Belém, Brazil
UFRJ Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil
USNM National Museum of Natural History, Washington, D.C., USA
Images and line drawings

Dorsal, ventral, and left lateral habitus images were taken using a DFC 450 C
Microsystems system (Leica, Wetzlar, Germany) with a Planapo 2.0x objective. Images
of subgenital plates, genitalia and forewings were taken with a Zeiss Axioskop 2
4 DOI 10.1163/1876312X-bja10029
compound microscope (Oberkochen, Germany) using Archimed V5.4.1 (Microvision).

The images were stacked using Zerene Stacker Vl.02 (Zerene Systems, Richland, WA,
USA).
To document the genitalia, the male and female abdomens were cleared after DNA
extraction using a Qiagen DNeasy® Blood & Tissue kit or a combination of the previ-
ous kit with a Qiagen QIAquick® PCR purification kit. Then, line drawings of cleared
genitalic features were made using a Nikon Eclipse 80i compound microscope (Nikon,
Tokyo, Japan) with a camera lucida.
Databasing
A matrix-code label with specimen identifier, consisting of a unique combination of an
eight-digit number and the prefix “UCR_ENT” was attached to each specimen, and
databased using the PBI instance of the Arthropod Easy Capture database served from
the AMNH https://research.amnh.org/pbi/locality/.
Maps
The distributional maps were created using QGIS v3.14 (https://www.qgis.org/) by
importing coordinates of specimens from the PBI database and from the specimen
labels.
Terminology
Terminology and abbreviations follow Leon & Weirauch (2016b) for general mor-
phology, Knyshov et al. (2020) for wing venation and Knyshov et al. (2018) for male
genitalic features.
Results
Taxonomy
Schizoptera (Zygophleps) McAtee and Malloch, 1925
Schizoptera (Zygophleps) McAtee and Malloch, 1925: 67, 18 (orig. descr.).
Type species: Schizoptera (Zygophleps) unica McAtee and Malloch, 1925, by original
designation
Diagnosis. Distinguished from other Schizopteridae by the posterolateral spine on
the metepisternum, the glabrous ventral margin of the metepisternum, the trapezoidal
shape of the discal cell of the forewing (Fig. 1A), the asymmetrical subgenital plate
(sternum 7) (Fig. 1G–H), the 90º rotation of the pygophore to the right of the dorsal
body plane, and the presence of two asymmetrical conjunctival appendages in males
(Fig. 1D–E). Distinguished from all other Schizoptera subgenera by the triangular
shape of the posterior membranal cell (dc1) of the forewing (Fig. 1A, black ellipse).
Re-description. Male. Body length ranging from 1.15 to 1.58 mm, body shape
ovoid, macropterous (Figs 2–10). COLORATION: Antennae yellow; head and thorax
DOI 10.1163/1876312X-bja10029 5
brown to dark brown; pronotum and scutellum brown to dark brown; legs yellow;
forewings anteriorly brown to dark brown, posteriorly pale brown; Sc + C concolorous
with or slightly lighter than remainder of hemelytron; hindwings hyaline; abdomen
brown to dark brown; right paramere either entirely concolorous (Figure 1B) or pro-
gressively darker toward apex. SURFACE and VESTITURE: Head and thorax with
dense cover of microtrichia; head, pronotum and wing veins with short to medium-
length light brown setae; clypeus and labium without microtrichia and with long setae;
numerous rounded muscle scars on frons; abdomen densely covered with long setae.
STRUCTURE: HEAD: flat or rounded in lateral view (Figs 8–10); ocelli roughly
equal in size to one ommatidium, positioned close to the margin of compound eye;
labium 3-segmented, last segment tapering. THORAX: pronotal collar pronounced,
approximately 1/7 length of pronotum; pronotum trapezoidal; scutellum triangular,
tip rounded; hemelytron distinctly punctate; costa+subcosta (C+Sc) enlarged; posterior
costal cell (rc1) squared; margins of wing veins outlined with small areoles; apex of wing
membranous with distinct cell-like sculpture; posterior membranal cell triangular (dc1,
Fig. 1A); metepisternum sculpted into a spine; ventral margin of metepisternum dif-
ferentiated into a narrow, glabrous shining groove; metasternal spine robust, elongate
and tapering to a point; hind coxa with well-developed adhesive pad; tarsal formula 3-3-
3; pretarsus with setiform parempodia; fore- and mid-tarsi with arolia. ABDOMEN:
with five visible sterna corresponding to segments 2 + 3, 4, 5, 6, and 7; subgenital plate
(enlarged sternum 7, st7) asymmetrical, modified into a subgenital plate with a single
lateral process on left side, posterior margin rounded or blunt (Figs 1G–H); medi-
otergites of pregenital abdomen equally sclerotized; mediotergite 7 (mt7) rectangular;
mediotergite 8 rectangular (mt8), slightly asymmetrical; length of mt8 about 3/4 pf
the length of mt7; three pairs of spiracles on segments 6–8. GENITALIA: pygophore
(py) rotated 90º to the right, open dorsally; parameres asymmetrical; right paramere
(rp) sickle shaped, with a robust base and elongate stem (Fig. 1B); left paramere (lp)
smaller than right, roughly quadrate, with two lobe-like processes (Fig. 1C); right con-
junctival appendage (rca) with base enlarged, flattened laterally, slightly tapering toward
apex, curved, branched or unbranched, with or without spines (Fig. 1D); left conjunc-
tival appendage (lca) smaller than right, with two arm-like projections of different sizes
(except in S. aculeata sp.n., S. cassiae sp.n., S. inaequalis sp.n., and S. magnifica sp.n.)
(Fig. 1E); vesica (v) coiled, forming four to six coils, tubular, thin, slightly tapering
toward apex (Fig. 1F); anophore (ano) well sclerotized; anal tube (at) membranous.
Female. Body length ranging from 1.23 to 1.51 mm, body shape ovoid, macropter-
ous (Fig. 11A–C). COLORATION and SURFACE and VESTITURE. As in male.
STRUCTURE. As in male, except by the tarsal formula 2-2-3, arolia absent; ster-
num 7 symmetrical, longer than preceding sterna, (Fig. 11E). GENITALIA. sternum
7 symmetrical; ovipositor vestigial; spermathecal gland (spgl) small and ellipsoidal;
spermathecal gland duct (spgd) straight; spermathecal reservoir (spr) bean-shaped, nar-
rower at the base and wider at apex; spermathecal duct (spd) long, thick, narrowing at
apex, coiled, forming five to six coils (Fig. 11H).
Distribution. Bolivia, Brazil, Costa Rica, Ecuador, French Guiana, Guatemala,
Peru, Suriname, and Trinidad and Tobago.
6 DOI 10.1163/1876312X-bja10029
Fig. 1. Diagnostic features of S. (Zygophleps) spp. A. Forewing (black ellipse showing the triangular shape
of the posterior membranal cell). B–J. Male. B. Right paramere (black arrow showing the median groove).
C. Left paramere. D. Right conjunctival appendage. E. Left conjunctival appendage (black arrows show-
ing the arm-like projections). F. Aedeagus. G–H Sternum 6 and the subgenital plate (sternum 7) (black
circle showing the posterolateral process of sternum 6). I–J Detail of the posterolateral process of sternum
6. Abbreviations: An1 = first anal vein, bc = basal cell, C = costa, Cu = cubitus, dc = discal cell, M = media,
R1 = first radial vein, R2 = second radial vein, rc1 = first radial cell, Sc = subcosta, scc = sub costal cell, tc
= trapezoidal cell.
Key to the species of Schizoptera (Zygophleps) based on males

This key includes all new species herein described and three species previously known
for which genitalic features were able to be assessed. Given that we were not able to
assess the male genitalia of S. unica McAtee & Malloch, this species was not included
here.
1. Posterior margin of the subgenital plate blunt (Figs 12, 13, e.g., S. applanata
sp.n., S. erosa sp.n., S. inaequalis sp.n.) ............................................................. 2
– Posterior margin of the subgenital plate rounded (Figs 13, 14 e.g., S. furcata sp.n.,
S. magnifica sp.n., S. priva sp.n.) ..................................................................... 12
DOI 10.1163/1876312X-bja10029 7
2. Right conjunctival appendage branched (Figs 20, 22 e.g., S. aculeata sp.n., S.

ramosa sp.n.) .................................................................................................... 3
– Right conjunctival appendage unbranched (Figs 20, 21 e.g., S. erosa sp.n., S. mal-
lochi sp.n., S. peruviana sp.n.) ........................................................................... 5
3. Process of the subgenital plate claw-shaped (Fig. 14) .................... S. ramosa sp.n.
– Process of the subgenital plate needle-shaped (Figs 12, 13, S. aculeata sp.n.,
S. inaequalis sp.n.) ............................................................................................ 4
4. Branches of the right conjunctival appendage of same length (Fig. 20) ................
................................................................................................. S. aculeata sp.n.
– Branches of the right conjunctival appendage of different lengths (Fig. 21) ..........
................................................................................................ S. inaequalis sp.n.
5. Process of the subgenital plate very short, little differentiated (Fig. 13, S. erosa
sp.n., S. kayi sp.n.) ........................................................................................... 6
– Process of the subgenital plate elongated, well-differentiated (Figs 13, 14, e.g., S.
fuscodorsata sp.n., S. mallochi sp.n., S. peruviana sp.n.) ..................................... 7
6. Right conjunctival appendage with a serrated margin (Fig. 20) ......... S. erosa sp.n.
– Right conjunctival appendage without serrated margin (Fig. 21) ....... S. kayi sp.n.
7. Process of the subgenital plate roughly rectangular (Fig. 14); right conjunctival
appendage with long spine on margin (Fig. 21) ............................. S. peruviana
sp.n.
– Process of subgenital plate “axe blade”-shaped (Fig. 12, S. applanata sp.n.), bean-
shaped (Fig. 13, S. fuscodorsata sp.n.) or thumb-like (Figs 13, 14, S. hirsuta sp.n.,
S. simla Emsley); right conjunctival appendage without long spine on margin
(Figs 20, 21, e.g., S. applanata sp.n., S. fuscodorsata sp.n.) ................................. 8
8. Process of subgenital plate “axe blade”-shaped (Figs 12, 14, S. applanata sp.n., S.
mallochi sp.n.) .................................................................................................. 9
– Process of subgenital plate bean-shaped (Fig. 13, S. fuscodorsata sp.n.) or thumb-
like (Figs 13, 14, S. hirsuta sp.n., S. simla Emsley) ............................................ 10
9. Anterior margin of process of subgenital plate anteriorly directed (Fig. 12); ser-
rated margin of right conjunctival appendage composed of medium-sized spines
(Fig. 20) .................................................................................. S. applanata sp.n.
– Anterior margin of process of subgenital plate anterolaterally directed (Fig. 14);
serrated margin of right conjunctival appendage composed of large spines (Fig. 21)
................................................................................................. S. mallochi sp.n.
10. Process of subgenital plate bean-shaped (Fig. 13); right conjunctival appendage
with serrated margin (Fig. 21) .............................................. S. fuscodorsata sp.n.
– Process of subgenital plate thumb-like (Fig. 13 e.g., S. hirsuta sp.n.); right con-
junctival appendage without serrated margin (Fig. 21 e.g., S. hirsuta sp.n.) ...... 11
11. Right conjunctival appendage with carinated margin; surface with small spines on
distal half (Fig. 21) ...................................................................... S. hirsuta sp.n.
– Right conjunctival appendage without carinated margin; surface with small spines
on apex (Fig. 22) ......................................................................... S. simla Emsley
12. Right conjunctival appendage branched (Figs 21, 22, e.g., S. furcata sp.n.,
S. stricta sp.n.) ................................................................................................ 13
8 DOI 10.1163/1876312X-bja10029
– Right conjunctival appendage unbranched (Figs 21, 22, e.g., S. knyshovi sp.n.,
S. speira sp.n.) ................................................................................................. 15
13. Process of subgenital plate claw-shaped (Fig. 12) ...................... S. corallia Emsley
– Process of subgenital plate long and straight (Figs 13, 15, S. furcata sp.n., S. stricta
sp.n.) .............................................................................................................. 14
14. Branches of right conjunctival appendage of same length and thin (Fig. 21) ........
..................................................................................................... S. furcata sp.n.
– Branches of right conjunctival appendage of different lengths and thick (Fig. 22)
....................................................................................................... S. stricta sp.n.
15. Right conjunctival appendage without spines (except in some specimens of S. kny-
shovi sp.n. that can have a small median spine on the margin) (Figs 20, 21, e.g.,
S. costarriquenha sp.n., S. inca sp.n.) .............................................................. 16
– Right conjunctival appendage with spines (Figs 20, 22, e.g., S. biancae sp.n.,
S. speira sp.n.) ................................................................................................. 20
16. Process of subgenital plate not laterally expanded on apex; anterolaterally directed
with wide base (Fig. 13) ................................................................... S. inca sp.n.
– Process of subgenital plate laterally expanded on apex; laterally directed with
narrow base (Figs 13, 14, e.g., S. falcata sp.n., S. knyshovi sp.n., S. paraensis sp.n.)
........................................................................................................................ 17
17. Process of subgenital plate “axe blade”-shaped (Fig. 13, S. falcata sp.n.) or “fish-
tail” shaped (Fig. 13, S. knyshovi sp.n.) ............................................................ 18
– Process of the subgenital plate oblong (Fig. 12, S. costarriquenha sp.n.) or heart-
shaped (Fig. 14, S. paraensis sp.n.) ................................................................... 19
18. Process of subgenital plate “axe blade”-shaped (Fig. 13) ............... S. falcata sp.n.
– Process of subgenital plate “fishtail”-shaped (Fig. 13) ................. S. knyshovi sp.n.
19. Process of subgenital plate oblong (Fig. 12) ...................... S. costarriquenha sp.n.
– Process of subgenital plate heart-shaped (Fig. 14) ...................... S. paraensis sp.n.
20. Right conjunctival appendage coiled (Fig. 22) ............................... S. speira sp.n.
– Right conjunctival appendage not coiled (Figs 20, 21, e.g., S. biancae sp.n.,
S. mcateei sp.n.) ............................................................................................. 21
21. Right conjunctival appendage surface with areolate sculptures (Fig. 21) ...............
.................................................................................................... S. mcateei sp.n.
– Right conjunctival appendage surface without areolate sculptures (Fig. 20 e.g.,
S. biancae sp.n., S. erwini sp.n.) ...................................................................... 22
22. Right conjunctival appendage without spines on surface (Figs 20, 21, e.g., S. bian-
cae sp.n., S. orellanensis sp.n.) ......................................................................... 23
– Right conjunctival appendage with spines on surface (Figs 20, 21, 22, e.g., S. cus-
pida sp.n., S. erwini sp.n., S. familia sp.n., S. pinduca sp.n.) ............................ 28
23. Process of subgenital plate goblet-shaped (Figs 12, 14, S. biancae sp.n., S. pichin-
cha sp.n.) ........................................................................................................ 24
– Process of subgenital plate thumb-like (Fig. 14, S. orellanensis sp.n.), trapezoidal
(Fig. 14, S. prima sp.n.), rectangular (Fig. 14, S. priva sp.n.) or roughly quadrate
(Fig. 15, S. ultima Emsley), but not goblet-shaped ........................................... 25
DOI 10.1163/1876312X-bja10029 9
24. Posterior margin of process of subgenital plate small, not clearly distinguishable
(Fig. 12); serrated margin of right conjunctival appendage composed of medium-
sized spines (Fig. 20); Costa Rica (Fig. 23) ................................... S. biancae sp.n.
– Posterior margin of process of subgenital plate long, clearly distinguishable (Fig.
14); serrated margin of right conjunctival appendage composed of long spines
(Fig. 21); Ecuador (Fig. 23) ....................................................... S. pichincha sp.n.
25. Process of subgenital plate thumb-like or trapezoidal (Fig. 14, S. orellanensis sp.n.,
S. prima sp.n.) ................................................................................................ 26
– Process of subgenital plate rectangular or roughly quadrate (Figs 14, 15, S. priva
sp.n., S. ultima Emsley) .................................................................................. 27
26. Process of subgenital plate thumb-like (Fig. 14, black square) ...............................
.............................................................................................. S. orellanensis sp.n.
– Process of subgenital plate trapezoidal (Fig. 14) ............................. S. prima sp.n.
27. Process of subgenital plate rectangular with median concavity on lateral margin
(Fig. 14, black arrow), small (Fig. 14) .............................................. S. priva sp.n.
– Process of subgenital plate roughly quadrate without median concavity, large
(Fig. 15) .................................................................................... S. ultima Emsley
28. Right conjunctival appendage surface with short serrate lines (Figs 20, 22,
S. erwini sp.n., S. pinduca sp.n.) ..................................................................... 29
– Right conjunctival appendage surface without short serrate lines (Figs 21, 22, e.g.,
S. familia sp.n., S. magnifica sp.n., S. solitaria sp.n.) ....................................... 30
29. Process of subgenital plate roughly rectangular (Fig. 13) ............... S. erwini sp.n.
– Process of subgenital plate spatulated (Fig. 14) ............................ S. pinduca sp.n.
30. Right conjunctival appendage without serrated margin; cluster of spines present
solely at apex (Figs 20, 21, e.g., S. ecuadoriana sp.n., S. familia sp.n.) ............. 31
– Right conjunctival appendage with serrated margin; cluster of spines present along
the surface (Figs 20, 21, e.g., S. cuspida sp.n., S. magnifica sp.n.) ................... 33
31. Right conjunctival appendage with long median spine on margin (Fig. 21) .........
.................................................................................................... S. familia sp.n.
– Right conjunctival appendage without median spine on margin (Fig. 20, S. ecua-
doriana sp.n., S. elegantissima sp.n.) ............................................................... 32
32. Process of subgenital plate shorter, broader, with anterior margin less projected
than S. elegantissima sp.n. (Fig. 12) ..................................... S. ecuadoriana sp.n.
– Process of subgenital plate longer, narrower, with anterior margin more projected
than S. ecuadoriana sp.n. (Fig. 12) ....................................... S. elegantissima sp.n.
33. Process of subgenital plate acute on apex (Fig. 12) ...................... S. cuspida sp.n.
– Process of subgenital plate truncate on apex (Figs 12, 13, e.g., S. cassiae sp.n., S. emsleyi
sp.n., S. magnifica sp.n.) ...……………………………………………………34
34. Process of subgenital plate remarkably enlarged, wider than long (Figs 13, 14, S.
magnifica sp.n., S. solitaria sp.n.) .................................................................... 35
– Process of subgenital plate not enlarged, longer than wide or as long as wide (Fig.
12, e.g., S. cassiae sp.n., S. emsleyi sp.n.) .......................................................... 36
10 DOI 10.1163/1876312X-bja10029
–35. Right conjunctival appendage with spines along entire distal half (Fig. 21) ..........
................................................................................................. S. magnifica sp.n.
– Right conjunctival appendage with spines solely at apex (Fig. 22) ........ S. solitaria
sp.n.
36. Process of subgenital plate remarkably elongated, about three times longer than
wide (Fig. 12) ............................................................................... S. emsleyi sp.n.
– Process of subgenital plate not elongated, as long as wide (Fig. 12, e.g., S. bella
sp.n., S. cassiae sp.n.) ...................................................................................... 37
37. Process of subgenital plate with tooth-like projection on posterior margin (Fig.
12, black arrow, S. bella sp.n., S. confusa sp.n.) .............................................. 38
– Process of subgenital plate without tooth-like projection on posterior margin
(Fig. 12) ....................................................................................... S. cassiae sp.n.
38. Process of subgenital plate with 1 tooth-like projection at apex (Fig. 12) ..............
........................................................................................................ S. bella sp.n.
– Process of subgenital plate with 2 or 3 tooth-like projections at apex (Fig. 12) .....
.................................................................................................... S. confusa sp.n.
Schizoptera (Zygophleps) aculeata sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/1B5D437B-6D2C-4722-BB6F-680CBCD5C40E
Diagnosis. Distinguished among species of S. (Zygophleps) by the long and curved
needle-shaped process of the subgenital plate (Fig. 12), and the right conjunctival
appendage long, thin, and branched with branches of same length and armed with
spines on apex (Fig. 20).
Measurements: Holotype. Total length: 1.41 mm; pronotum length: 0.20 mm;
pronotum width: 0.56 mm; scutellum length: 0.15 mm; abdominal width: 0.53 mm.
Material examined (n = 5): HOLOTYPE: ECUADOR: Orellana: Tiputini
Biodiversity Station Co.: nr Yasuni National Park Erwin Transect - T/4, 0.63194°N
76.14417°W, 220 m, 04 Oct 1996, T. L. Erwin et al., 1♂ (UCR_ENT 00029060)
(USNM). PARATYPES: ECUADOR: Orellana: Same data as holotype, 1♂ (UCR_
ENT 00028839) (USNM); Transect - T/4, 24 Oct 1998, T. L. Erwin et al., 1♂ (UCR_
ENT 00028305) (USNM); Transect - T/9, 0.63194°N 76.14417°W, 220 m, 05 Feb
1999, T. L. Erwin et al., 1♂ (UCR_ENT 00028811) (USNM). Transect Ent. 1 km
S. Onkone Gare Camp, Reserva Etnica Waorani, Onkone Gare Camp, 0.65714°N
(USNM).
Description. Male (Figs 2, 5, 8). COLORATION: General coloration brown; C+Sc
lighter than remainder of hemelytra (Fig. 2); rp almost entirely pale brown with apex
brown. SURFACE and VESTITURE: as in subgeneric description. STRUCTURE:
HEAD: rounded in lateral view (Fig. 8). THORAX: as in subgeneric description.
ABDOMEN: as in subgeneric description; sternum 6 asymmetrical with a medium-
sized posterolateral triangular process slightly projecting on the anterolateral margin
DOI 10.1163/1876312X-bja10029 11
of subgenital plate (Fig. 12); subgenital plate weakly asymmetrical, posterior margin
blunt (Fig. 12). GENITALIA: py as in subgeneric description; rp large, with a deep
median groove, flattened dorsoventrally, apex with two lateral projections (Fig. 16);
lp as in subgeneric description; lca with one arm-like projection; v, ano, and at as in
subgeneric description.
Female. Unknown.
Distribution (Fig. 23). ECUADOR: Orellana.
Etymology. Named for the needle-shaped process of the subgenital plate after the
Latin “aculeata” meaning needle.
Comments. This species resembles S. inaequalis sp.n. in the shape of the subgenital
plate and its process, however the two species differ in the shape of the branches of
the right conjunctival appendages. While S. aculeata sp.n. has branches of the same
length, S. inaequalis sp.n. has branches of different lengths.
Schizoptera (Zygophleps) applanata sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/BC51A94F-1366-4FD7-A324-A7B25F8B4331
Diagnosis. Distinguished among species of S. (Zygophleps) by the strongly asymmetri-
cal subgenital plate with posterior margin remarkably blunt (Fig. 12), the “axe blade”-
shaped process of the subgenital plate with anterior margin pointed anteriorly and
elongated and posterior margin rounded and short (Fig. 12), and the long, robust, and
unbranched right conjunctival appendage with a serrated margin (Fig. 20).
Biodiversity Station, nr Yasuni National Park, Erwin Transect - T/8, 0.63194°N
(USNM). PARATYPES: ECUADOR: Napo: Waorani, 1 km S. Onkone Gare
Camp, 0.63333°N 76.6°E, 220 m, 21 Jun 1994, T. L. Erwin et al., 1♂ (UCR_ENT
00026398) (USNM); Orellana: Tiputini Biodiversity Station, nr Yasuni National
Park, Erwin Transect - T/8, 0.63194°N 76.14417°W, 220 m, 06 Feb 1999, T. L. Erwin
et al., 1♂ (UCR_ENT 00028546) (USNM); Same data as holotype, 1♂ (UCR_ENT
00128276) (USNM). Transect - T/10, 0.63194°N 76.14417°W, 9♂ (UCR_ENT
00027852, UCR_ENT 00029145, UCR_ENT 00029146, UCR_ENT 00029147,
UCR_ENT 00029148, UCR_ENT 00029149, UCR_ENT 00029150, UCR_ENT
00029151, UCR_ENT 00029152) (USNM).
Description. Male (Figs 2, 5, 8). COLORATION: General coloration brown;
C+Sc slightly lighter than remainder of hemelytra (Fig. 2); rp entirely dark brown.
SURFACE and VESTITURE: as in subgeneric description. STRUCTURE: HEAD:
flat in lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as in
subgeneric description; sternum 6 strongly asymmetrical with a large triangular poste-
rolateral process projecting from the anterolateral margin of subgenital plate (Fig. 12).
12 DOI 10.1163/1876312X-bja10029
GENITALIA: py as in subgeneric description; rp large, with a deep median groove,

apex long, curved and pointed (Fig. 16); lp as in subgeneric description; lca with the
large projection about two times longer than the small; v, ano, and at as in subgeneric
description.
Female. Unknown.
Distribution (Fig. 23). ECUADOR: Napo, Orellana.
Etymology. Named for the laterally flattened right conjunctival appendage after the
Latin “applanatus” meaning flattened.
Comments. This species resembles S. mallochi sp.n. in having similar processes of
the subgenital plate and right conjunctival appendage. However, there are some subtle
differences between the two species. The anterior margin of the process of the sub-
genital plate in S. applanata sp.n. is anteriorly directed while in S. mallochi sp.n. this
margin is anterolaterally directed. In addition, the spines on the serrated margin of the
right conjunctival appendage in S. applanata sp.n. are smaller than in S. mallochi sp.n.
Schizoptera (Zygophleps) bella sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/19BF5C5D-3953-4D4B-B81F-439407A3525B
Diagnosis. Distinguished among species of S. (Zygophleps) by the roughly quadrate
process of the subgenital plate with posterior margin elongated, pointed and laterally
directed (Fig. 12, black arrow), and the small, robust, and unbranched right conjuncti-
val appendage with serrated margin and surface with small spines on the median third
(Fig. 20).
Material examined (n = 8): HOLOTYPE: ECUADOR: Orellana: Estacion Cientifica
Yasuni, 0.67444°N 76.64722°W, 215 m, 05 Sep 1999 - 10 Sep 1999, E.G. Riley,
1♂ (UCR_ENT 00093586) (TAMU). PARATYPES: BRAZIL: Rondônia: Fazenda
Rancho Grande, 62 km S. Ariquenes, 05 Jul 1991, S.L. Heydon, 1♂ (UCR_ENT
00037078) (UCD). ECUADOR: Orellana: Tiputini Biodiversity Station near Yasuni
National Park Erwin Transect - T/6, 0.63194°N 76.14417°E, 220 m, 26 Oct 1998,
T. L. Erwin et al., 2♂ (UCR_ENT 00028276, UCR_ENT 00028902) (USNM);
Transect - T/10, 21 Oct 1998, T. L. Erwin et al., 1♂ (UCR_ENT 00028205) (USNM);
04 Oct 1996, T. L. Erwin et al., 2♂ (USNM) (UCR_ENT 00028840, UCR_ENT
00029062) (USNM); 05 Feb 1999, T. L. Erwin et al., 1♂ (UCR_ENT 00027822)
(USNM).
lighter than remainder of hemelytra (Fig. 2); rp entirely pale brown. SURFACE and
VESTITURE: as in subgeneric description. STRUCTURE: HEAD: slightly rounded
in lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as in
subgeneric description; sternum 6 weakly asymmetrical with a small rounded pos-
terolateral process slightly projecting from the anterior margin of subgenital plate
DOI 10.1163/1876312X-bja10029 13
(Fig. 12); subgenital plate weakly asymmetrical, posterior margin rounded (Fig. 12).
GENITALIA: py as in subgeneric description; rp small, with a shallow median groove,
apex short, slightly curved and rounded (Fig. 16); lp as in subgeneric description; lca
with the large projection about two times longer than the small; v, ano, and at as in
Female. Unknown.
Distribution (Fig. 23). BRAZIL: Rondônia; ECUADOR: Orellana.
Etymology. Named for the beauty of this species after the Latin “bellus” meaning
beautiful.
Comments. The process of the subgenital plate distinguishes this species from all
other species in this subgenus.
Schizoptera (Zygophleps) biancae sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/B0BF4C68-5981-4970-ABF4-BF5EFE69C35A
Diagnosis. Distinguished among species of S. (Zygophleps) by the process of the sub-
genital plate with anterior margin well-developed and rounded and posterior margin
small and pointed (Fig. 12), and the long, robust, and unbranched right conjunctival
appendage with a serrated margin (Fig. 20).
Material examined (n = 3): HOLOTYPE: COSTA RICA: Heredia: La Selva
Biological Station, 10.43333°N 84.01667°W, 56 m, 05 Jan 2000, Eugenia sp.
(Myrtaceae), 1♂ (UCR_ENT 00109292) (INBIO). PARATYPES: COSTA
RICA: Heredia: La Selva Biological Station, 10.43333°N 84.01667°W, 56 m, 02
Feb 1998, ALAS, Light Trap, 2♂ (UCR_ENT 00109228, UCR_ENT 00109229)
(INBIO).
C+Sc concolorous with remainder of hemelytra (Fig. 2); rp brown with apex dark
ABDOMEN: as in subgeneric description; sternum 6 weakly asymmetrical with a
small rounded posterolateral process slightly projecting from the anterolateral margin
rounded (Fig. 12). GENITALIA: py as in subgeneric description; rp small, without a
median groove, folded, apex pointed and laterally directed in dorsal view [not shown
in the drawing] (Fig. 16); lp as in subgeneric description; lca with the large projection
about three times longer than the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Distribution (Fig. 23). COSTA RICA: Heredia.
Etymology. Named as a homage to Bianca Albuquerque.
14 DOI 10.1163/1876312X-bja10029
Comments. This species shares with S. pichincha sp.n. the unique shape of the right
paramere. However, the two species can be distinguished based on the shape of the
process of the subgenital plate and the right conjunctival appendage.
Schizoptera (Zygophleps) cassiae sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/B31DEA32-B713-4153-B51F-809C50114659
Diagnosis. Distinguished among species of S. (Zygophleps) by the rounded process of
the subgenital plate with a median concavity and a rounded anterior margin antero-
laterally directed (Fig. 12), and the long, robust, and unbranched right conjunctival
appendage with surface composed of medium-sized spines on the second half (Fig. 20).
Measurements: Total length: 1.31 mm; pronotum length: 0.24 mm; pronotum
width: 0.61 mm; scutellum length: 0.13 mm; abdominal width: 0.55 mm.
Biological Station, 10.43333°N 84.01667°W, 56 m, 02 Feb 1998, ALAS, Light Trap,
1♂ (UCR_ENT 00109230) (INBIO).
slightly lighter than remainder of hemelytra (Fig. 2); rp entirely pale brown. SURFACE
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: rounded in
lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as in sub-
generic description; sternum 6 weakly asymmetrical with a small rounded posterolat-
eral process slightly projecting from the anterolateral margin of the subgenital plate
GENITALIA: py as in subgeneric description; rp large, with a strongly deep median
groove, apex long, curved and rounded (Fig. 16); lp as in subgeneric description; lca
with arm-like projections of about the same size, one projection dorsally and another
ventrally directed; v, ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named as a homage to Cássia Pereira.
Comments. The process of the subgenital plate of this species is an excellent diag-
nostic character to distinguish it from other species of S. (Zygophleps).
Schizoptera (Zygophleps) confusa sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/B9631B6A-8362-43B5-A308-3EB2C76B4537
Diagnosis. Distinguished among species of S. (Zygophleps) by the roughly rectangular
process of the subgenital plate with one small tooth-like projection on each margin and
one median small tooth-like projection, which can be absent in some specimens (Fig.
12), and the long, thin, and unbranched right conjunctival appendage with surface
composed of small spines on the second half (Fig. 20).
DOI 10.1163/1876312X-bja10029 15

Biological Station, 10.43333°N 84.01667°W, 56 m, 05 Jan 2000, Eugenia sp.
(Myrtaceae), 1♂ (UCR_ENT 00109294) (INBIO). PARATYPES: COSTA
RICA: Heredia: Same data as holotype 1♂ (UCR_ENT 00109295) (INBIO); 10
Oct 1994, ALAS, Ilex sp. (Aquifoliaceae), (UCR_ENT 00109103) 1♂ (INBIO); 16
Apr 1998, Light Trap, 1♂ (UCR_ENT 00109214) (INBIO); 02 Jun 1998, Light
Trap, 1♂ (UCR_ENT 00109191) (INBIO). PERU: Loreto: Maynas, Calliceb
us Res. Station Mishana, Rio Nanay, 25km SW Iquitos, 3.90888°S 73.41394°W,
120 m, 10–17 Jan 1980, J. B. Heppner, 1♂ (UCR_ENT 00026412) (USNM).
TRINIDAD AND TOBAGO: Arima: Simla, Simla Research Station, off verand at
station, 10.6928°N 61.2897°W, 250 m, 22 Jul 2013, C. Weirauch, 1♂ (UCR_ENT
00102132) (UCR); Belmont: 10 km NE Roxborough, Gilpin Tr., 11.31671°N
60.529°W, 400 m, 26–30 Nov 1993, S. B. Peck, J. Kukalova-Peck, 1♂ (UCR_ENT
00091204) (FMNH).
C+Sc slightly lighter than remainder of hemelytra (Fig. 2); rp entirely pale brown.
rounded in lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN:
as in subgeneric description; sternum 6 asymmetrical with a medium-sized triangular
posterolateral process strongly projecting from the anterolateral margin of subgeni-
tal plate (Fig. 12); subgenital plate weakly asymmetrical, posterior margin rounded
(Fig. 12). GENITALIA: py as in subgeneric description; rp small, with a deep median
groove, apex short, slightly curved and pointed (Fig. 16); lp as in subgeneric descrip-
tion; lca with the large projection about two times longer than the small; v, ano, and
at as in subgeneric description.
Female. Unknown.
Distribution (Fig. 23). COSTA RICA: Heredia; PERU: Loreto; TRINIDAD AND
TOBAGO: Arima, Belmont.
Etymology. Named for the morphological variation of the process of the subgenital
plate, which made it difficult to determine the concept of this species.
Comments. This species shows morphological variability of the process of the sub-
genital plate. The median tooth-like projection on the apex of this process is present in
some specimens and absent in others.
Schizoptera (Zygophleps) corallia Emsley, 1969

Schizoptera (Zygophleps) corallia Emsley, 1969: 25, 67 (orig. descr.)
(Figs 12, 16, 20, 23)
Updated diagnosis. Distinguished among species of S. (Zygophleps) by the claw-
shaped process of the subgenital plate posteriorly directed (Fig. 12), the right paramere
with a deep median groove, apex short, slightly curved and pointed [missing in the
16 DOI 10.1163/1876312X-bja10029
holotype] (Fig. 16, adapted from Emsley (1969)), and the long, thin, roughly straight,
and branched right conjunctival appendage with branches of about same length and
armed with spines on apex (Fig. 20).
Measurements: Total length: 1.50 mm; pronotum length:? mm; pronotum width:?
mm; scutellum length: 0.15 mm; abdominal width: 0.60 mm [question marks (?) stand
for body parts that could not be measured given their position on the slide].
Material examined (n = 1): HOLOTYPE: TRINIDAD AND TOBAGO: Arima
Valley: William Beebe Tropical Research Station, at light, 29 Sep 1961, M. G. Emsley,
1♂ (AMNH_IZC 00150327) (AMNH).
Re-description. Male. COLORATION: General coloration brown; C+Sc con-
colorous with remainder of hemelytra. SURFACE and VESTITURE: as in subgeneric
description. STRUCTURE: THORAX: as in subgeneric description. ABDOMEN:
as in subgeneric description; sternum 6 weakly asymmetrical with a medium-sized
posterolateral triangular process projecting from the anterolateral margin of subgenital
plate (Fig. 12); subgenital plate weakly asymmetrical, posterior margin rounded (Fig.
12). GENITALIA: py as in subgeneric description; lp as in subgeneric description; lca
Female. Unknown.
Distribution (Fig. 23). TRINIDAD AND TOBAGO: Arima Valley.
Comments. Emsley (1969) described this species from Trinidad based on a single
male specimen. He mentioned the distinct shape of the process of the subgenital plate
and right conjunctival appendage among species of S. (Zygophleps). These two features
set apart S. corallia from all other species of this subgenus, except for S. ramosa sp.n.,
and we initially suspected that specimens assigned to the two species may be conspe-
cific. However, while the right paramere of the holotype of S. corallia is missing and
we were unable to examine it directly, the drawing by Emsley (reproduced here in Fig.
16) shows that it is very different from the right paramere in S. ramosa sp.n. For this
reason, we consider the two as different species.
Schizoptera (Zygophleps) costarriquenha sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/FDD35509-5654-43D7-A484-447F7AA654FD
Diagnosis. Distinguished among species of S. (Zygophleps) by the oblong process of
subgenital plate process of the subgenital plate (Fig. 12), and the small, thin, and
unbranched right conjunctival appendage without spines (Fig. 20).
Biological Station, 10.43333°N 84.01667°W, 56 m, 05 Jan 2000, ALAS, Eugenia sp.
(Myrtaceae), 1♂ (UCR_ENT 00109293) (INBIO).
DOI 10.1163/1876312X-bja10029 17

concolorous with remainder of hemelytra (Fig. 2); rp entirely pale brown. SURFACE
generic description; sternum 6 weakly asymmetrical without process (Fig. 12); sub-
genital plate weakly asymmetrical, posterior margin rounded (Fig. 12). GENITALIA:
py as in subgeneric description; rp medium-sized, with a deep median groove, apex
medium, slightly curved and rounded (Fig. 16); lp as in subgeneric description; lca
Female. Unknown.
Etymology. Named after the country of origin of this species after the word “costar-
riquenha” meaning from Costa Rica.
Comments. The process of the subgenital plate of this species is somewhat similar
to S. fuscodorsata sp.n., but the anterior margin is more laterally projected in S. costar-
riquenha sp.n. Additionally, the shape of the right conjunctival appendages in the two
species is distinct. The margin of the right conjunctival appendage is not serrated in S.
costarriquenha sp.n., while it is serrated in S. fuscodorsata sp.n.
Schizoptera (Zygophleps) cuspida sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/F19DDAAF-EB8F-4851-A3CA-586AF1498E9A
Diagnosis. Distinguished among species of S. (Zygophleps) by the long, straight, taper-
ing, remarkably acute on apex process of the subgenital plate (Fig.12), and the long,
thin, and unbranched right conjunctival appendage with a serrated margin and surface
with small spines on the second third (Fig. 20).
Biodiversity Station Co.: nr Yasuni National Park Erwin Transect - T/10, 0.63194°N
(USNM). PARATYES: ECUADOR: Orellana: Res. Ethnica Waorani, 1km S.
Onkone Gare Camp, 0.63333°N 76.6°E, 220 m, 21 Jun 1994, T. L. Erwin et al.,
1♂ (UCR_ENT 00030648) (USNM); Tiputini Biodiversity Station Co.: nr Yasuni
National Park Erwin Transect - T/4, 0.63194°N 76.14417°W, 220 m, 24 Oct 1998,
T. L. Erwin et al., 1♂ (UCR_ENT 00028528) (USNM).
as in subgeneric description; sternum 6 weakly asymmetrical with a small rounded
18 DOI 10.1163/1876312X-bja10029
posterolateral process slightly projecting from the anterolateral margin of subgenital

12). GENITALIA: py as in subgeneric description; rp small, with a shallow median
groove, apex medium, slightly curved and rounded (Fig. 16); lp as in subgeneric
description; lca with the large projection about three times longer than the small; v,
ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named for the pointed process of the subgenital plate after the Latin
“cuspis” meaning pointed end.
Comments. The pointed and laterally directed process of the subgenital plate is a
distinctive character among other species of this subgenus.
Schizoptera (Zygophleps) ecuadoriana sp.n.

(Figs 2, 5, 8, 12, 16, 20, 23)
ZooBank: http://zoobank.org/58156ECE-72D7-4C07-B215-89B8DF73F469
Diagnosis. Distinguished among species of S. (Zygophleps) by the roughly triangular
process of the subgenital plate (Fig. 12), and the small, thin, and unbranched right
conjunctival appendage with small spines on apex (Fig. 20).
Biodiversity Station nr Yasuni National Park; Erwin transect T/1, 220-250m, 0.63194°S
76.14417°W, 220 m, 09 Nov 1999, T. L. Erwin et al., 1♂ (UCR_ENT 00027919)
(USNM). PARATYPES: ECUADOR: Orellana: Transect Ent. 1 km S. Onkone Gare
Camp, Reserva Etnica Waorani, Onkone Gare Camp, 0.65714°N 76.453°W, 216 m,
04 Oct 1996, T. L. Erwin et al., 3♂ (UCR_ENT 00028841, UCR_ENT 00028842,
UCR_ENT 00029061) (USNM).
slightly concolorous with remainder of hemelytra (Fig. 2); rp entirely pale brown.
flat in lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as
in subgeneric description; sternum 6 weakly asymmetrical with a small rounded pos-
terolateral process slightly projecting from the anterolateral margin of subgenital plate
apex medium, slightly curved and rounded (Fig. 16); lp as in subgeneric description;
lca with the large projection about two times longer than the small; v, ano, and at as in
Female. Unknown.
DOI 10.1163/1876312X-bja10029 19
Etymology. Named after the country where the species was collected after the word
“ecuadoriana” meaning from Ecuador.
Comments. The triangular process of the subgenital plate is unique among species
of this subgenus.
Schizoptera (Zygophleps) elegantissima sp.n.

(Figs 2, 5, 8, 12, 17, 20, 23)
ZooBank: http://zoobank.org/6DDD9EA8-766B-4B2E-BE30-AE6F0372CE45
Diagnosis. Distinguished among species of S. (Zygophleps) by the “shark tail”-shaped
process of the subgenital plate (Fig. 12), and the small, thin, and unbranched right
conjunctival appendage with small spines on apex (Fig. 20).
Material examined (n = 1): HOLOTYPE: ECUADOR: Orellana: Transect Ent.
1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 0.65714°N 76.453°W, 216 m,
02 Oct 1996, T. L. Erwin et al., 1♂ (UCR_ENT 00027873) (USNM).
C+Sc concolorous with remainder of hemelytra (Fig. 2); rp entirely pale brown.
flat in lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as
in subgeneric description; sternum 6 weakly asymmetrical with a medium-sized pos-
terolateral process slightly projecting from the anterolateral margin of subgenital plate
apex short, slightly curved and pointed (Fig. 17); lca with the large projection about
two times longer than the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named for the beauty of this species after the word “elegantissima”
meaning very beautiful.
Comments. This species ressembles S. ecuadoriana sp.n. by the the process of sub-
genital plate. However, this process in S. elegantissima sp.n. is longer, narrower, with
anterior margin more projected than S. ecuadoriana sp.n.
Schizoptera (Zygophleps) emsleyi sp.n.

(Figs 2, 5, 8, 12, 17, 20, 23)
ZooBank: http://zoobank.org/C074CAF1-0F8C-4C63-8E71-3A2D5F2D35C6
Diagnosis. Distinguished among species of S. (Zygophleps) by the thin and elongated
process of the subgenital plate with anterior margin projecting anterolaterally (Fig.
12), and the long, thin, and unbranched right conjunctival appendage with a serrated
margin and surface with small spines on the second half (Figure 20).
20 DOI 10.1163/1876312X-bja10029

Material examined (n = 1): HOLOTYPE: ECUADOR: Orellana: Waorani, 1km
S. Onkone Gare Camp, 0.63333°N 76.6°W, 220 m, 25 Jun 1994, T. L. Erwin et al.,
1♂ (UCR_ENT 00026410) (USNM).
lighter than remainder of hemelytra (Fig. 2); rp pale brown with apex brown. SURFACE
generic description; sternum 6 strongly asymmetrical with a large posterolateral process
projecting from the anterolateral margin of subgenital plate and reaching its process
GENITALIA: py as in subgeneric description; rp large, with a shallow median groove,
apex extremely long, thin, slightly curved and pointed (Fig. 17); lp as in subgeneric
description; lca with the large projection about two times longer than the small; v, ano,
and at as in subgeneric description.
Female. Unknown.
Etymology. Named in Honor of Dr. Michael Emsley, who made important con-
tributions to the knowledge of Dipsocoromorpha and described three species of S.
(Zygophleps).
Comments. This species shares the similar shape of the process of the subgenital plate
with S. speira sp.n. Despite this similarity, these species differ in the shape of the right
conjunctival appendages.
Schizoptera (Zygophleps) erosa sp.n.

(Figs 2, 5, 8, 13, 17, 20, 23)
ZooBank: http://zoobank.org/3EE7ACF1-8816-4B4A-BC02-53C7A8324D5A
Diagnosis. Distinguished among species of S. (Zygophleps) by the very short and
roughly quadrate process of the subgenital plate with apex slightly concave and with
anterior margin rounded and posterior margin slightly pointed (Fig. 13), and the long,
thin, and unbranched right conjunctival appendage with a serrated margin (Fig. 20).
Material examined (n = 2): HOLOTYPE: ECUADOR: Orellana: Transect Ent.
1 km S. Onkone Gare Camp, Reserva Etnica Waorani, Onkone Gare Camp,
0.65714°N 76.453°W, 216 m, 03 Oct 1996, T. L. Erwin et al., 1♂ (UCR_ENT
00028081) (USNM). PARATYPE: ECUADOR: Orellana: Tiputini Biodiversity
Station nr Yasuni National Park Erwin Transect - T/10, 0.63194°N 76.14417°W,
220 m, 21 Oct 1998, T. L. Erwin et al., 1♂ (UCR_ENT 00028206) (USNM).
Description. Male (Figes 2, 5, 8). COLORATION: General coloration brown;
DOI 10.1163/1876312X-bja10029 21

as in subgeneric description; sternum 6 weakly asymmetrical with a small rounded pos-
terolateral process projecting from the anterior margin of subgenital plate (Fig. 13);
subgenital plate weakly asymmetrical, posterior margin blunt (Fig. 13). GENITALIA:
py as in subgeneric description; rp large, with a shallow median groove, apex short,
curved and rounded (Fig. 17); lp as in subgeneric description; lca with the large projec-
tion about two times longer than the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named for the serrated right conjunctival appendage after the Latin
“erosus” meaning serrated.
Comments. The process of the subgenital plate of S. erosa sp.n. is very short com-
pared to other S. (Zygophleps) species and resembles S. kayi sp.n. Despite this similar-
ity, these species can be distinguished from each other based on the shape of the right
conjunctival appendage.
Schizoptera (Zygophleps) erwini sp.n.

(Figs 2, 5, 8, 13, 17, 20, 23)
ZooBank: http://zoobank.org/11CD8978-0DD8-4A58-9479-A0CBE9280780
process of the subgenital plate with anterior and posterior margins rounded (Fig. 13),
and the small, thin, and unbranched right conjunctival appendage with a serrated
margin composed of small spines and with short serrate lines on the surface (Fig. 20).
Biodiversity Station nr Yasuni National Park Erwin Transect - T/10, 0.63194°N
(USNM). PARATYPE: ECUADOR: Orellana: Transect Ent. 1 km S. Onkone Gare
Camp, Reserva Etnica Waorani, Onkone Gare Camp, 0.65714°N 76.453°W, 216 m,
02 Oct 1996, T. L. Erwin et al., 1♂ (UCR_ENT 00027863) (USNM).
Description. Male (Figs 2, 5, 8). COLORATION: General coloration pale brown;
C+Sc lighter than remainder of hemelytra (Fig. 2); rp entirely pale brown. SURFACE
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: flat in lat-
eral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as in sub-
generic description; sternum 6 weakly asymmetrical with a thumb-like posterolateral
process laterad-directed slightly projecting from the anterolateral margin of the sub-
genital plate (Fig. 13); subgenital plate weakly asymmetrical, posterior margin rounded
(Fig. 13). GENITALIA: py as in subgeneric description; rp large, apex medium-sized,
22 DOI 10.1163/1876312X-bja10029
Female. Unknown.
Etymology. Named in Honor of Dr. Terry Erwin, who collected several species of
S. (Zygophleps) herein described.
Comments. This species resembles S. peruviana sp.n. and S. priva sp.n. in having
a similar process of the subgenital plate. However, these species can be easily distin-
guished from each other based on the shape of the right conjunctival appendage.
Schizoptera (Zygophleps) falcata sp.n.

(Figs 2, 5, 8, 13, 17, 20, 23)
ZooBank: http://zoobank.org/D4BAF678-F925-4E93-91B2-7890FC618EEB
Diagnosis. Distinguished among species of S. (Zygophleps) by the “axe blade”-shaped
process of the subgenital plate with anterior margin pointed and posterior margin
rounded (Fig. 13), and the long, robust, and unbranched right conjunctival appendage
without spines (Fig. 20).
Biodiversity Station nr Yasuni National Park Erwin Transect - T/8, 0.63194N,
-76.14417W, 220 m, 06 Feb 1999, T. L. Erwin et al., canopy fogging, 1♂ (UCR_ENT
00028545) (USNM). PARATYPE: ECUADOR: Orellana: Tiputini Biodiversity
Station nr Yasuni National Park Erwin Transect - T/8, 0.63194N, -76.14417W, 220
m, 06 Feb 1999, T. L. Erwin et al., canopy fogging, 1♂ (UCR_ENT 00027890)
(USNM); Transect Ent. 1 km S. Onkone Gare Camp, Reserva Etnica Waorani,
Onkone Gare Camp, 0.65714°N 76.453°W, 03 Oct 1996, T. L. Erwin et al., 1♂
(USNM) (UCR_ENT 00028087) (USNM).
flat in lateral view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN:
as in subgeneric description; sternum 6 weakly asymmetrical with a small rounded
posterolateral process slightly projecting from the anterior margin of subgenital plate
(Fig. 13); subgenital plate asymmetrical, posterior margin slightly rounded (Fig. 13).
GENITALIA: py as in subgeneric description; rp large, with a shallow median groove,
apex extremely long, curved and rounded (Fig. 17); lp as in subgeneric description; lca
with the large projection about three times longer than the small; v, ano, and at as in
Female. Unknown.
Etymology. Named for the shape of the right paramere after the Latin “falcatus”
meaning sickle shaped.
DOI 10.1163/1876312X-bja10029 23
Comments. This species resembles S. applanata sp.n. and S. mallochi sp.n. in hav-
ing a similar process of the subgenital plate. Nevertheless, these species can be distin-
guished based on the shape of the right conjunctival appendage. The margin of the
right conjunctival appendage is not serrated in S. falcata sp.n., while it is serrated in S.
applanata sp.n. and S. mallochi sp.n.
Schizoptera (Zygophleps) familia sp.n.

(Figs 2, 5, 8, 13, 17, 21, 23)
ZooBank: http://zoobank.org/4DD0C1B1-7DE2-4F74-ACB9-23AEB1B058E7
Diagnosis. Distinguished among species of S. (Zygophleps) by the thumb-like process
of the subgenital plate (Fig. 13), and the long, thin, and unbranched right conjunc-
tival appendage with a long median spine on the margin and apex with small spines
(Fig. 21).
-76.14417W, 220 m, 26 Oct 1998, T. L. Erwin et al., canopy fogging, 1♂ (UCR_
ENT 00128274) (USNM).
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: flat in lateral
view (Fig. 8). THORAX: as in subgeneric description. ABDOMEN: as in subgeneric
description; sternum 6 weakly asymmetrical with a small rounded posterolateral process
slightly projecting from the anterolateral margin of subgenital plate (Fig. 13); subgeni-
tal plate weakly asymmetrical, posterior margin rounded (Fig. 13). GENITALIA: py
as in subgeneric description; rp small, with a deep median groove, apex long, strongly
Female. Unknown.
Etymology. Named as a homage to our families that are extremely important in our
lives after the Latin “familia” meaning family.
Comments. See comments on S. hirsuta sp.n. and S. simla Emsley.
Schizoptera (Zygophleps) furcata sp.n.

(Figs 2, 5, 8, 13, 17, 21, 23)
ZooBank: http://zoobank.org/2247B638-BC42-446C-BFD2-98BE6B931FB1
Diagnosis. Distinguished among species of S. (Zygophleps) by the long, straight and
anterolaterally directed process of the subgenital plate (Fig. 13), and the long, thin,
and branched right conjunctival appendage with branches of same length and without
spines (Fig. 21).
24 DOI 10.1163/1876312X-bja10029

-76.14417W, 220 m, 26 Oct 1998, T. L. Erwin et al., canopy fogging, 1♂ (UCR_
ENT 00028901) (USNM).
as in subgeneric description; sternum 6 weakly asymmetrical with a small posterolat-
eral process laterad-directed (Fig. 13); subgenital plate weakly asymmetrical, posterior
margin rounded (Fig. 13). GENITALIA: py as in subgeneric description; rp large, with
a very deep median groove, apex extremely long, strongly curved and rounded (Fig.
17); lp as in subgeneric description; lca with the large projection about three times
longer than the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named for the forked right conjunctival appendage after the Latin “fur-
catus” meaning forked.
Comments. This species can be easily recognized among species of S. (Zygophleps)
by the unique shape of the process of the subgenital plate. Additionally, the right con-
junctival appendage is unique within the subgenus.
Schizoptera (Zygophleps) fuscodorsata sp.n.

(Figs 3, 6, 9, 13, 17, 21, 23)
ZooBank: http://zoobank.org/3E69BDA7-E217-40D7-96AB-B9D023B314DE
Diagnosis. Distinguished among species of S. (Zygophleps) by the bean-shaped process
of the subgenital plate (Fig. 13), and the small, thin, and unbranched right conjunc-
tival appendage with a serrated margin composed of medium-sized spines (Fig. 21).
Material examined (n = 1): HOLOTYPE: COSTA RICA: Heredia: La Selva Biol.
Sta., 10.43056°N 84.00667°W, 58 m, 14 Aug 2010, J. Mottern, 1♂ (UCR_ENT
00081617) (UCR).
Description. Male (Figs 3, 6, 9). COLORATION: General coloration dark brown;
C+Sc concolorous with remainder of hemelytra (Fig. 3); rp brown with apex dark brown.
as in subgeneric description; sternum 6 asymmetrical with a medium-sized rounded
posterolateral process projecting from the anterolateral margin of subgenital plate
(Fig. 13); subgenital weakly strongly asymmetrical, posterior margin blunt (Fig. 13).
DOI 10.1163/1876312X-bja10029 25

apex long, short and pointed (Fig. 17); lp as in subgeneric description; lca with the
description.
Female. Unknown.
Etymology. Named for the dorsal dark brown coloration after the Latin “fuscus”
and “dorsum” meaning dark and back, respectively.
Comments. See comments on S. costarriquenha sp.n.
Schizoptera (Zygophleps) hirsuta sp.n.

(Figs 3, 6, 9, 13, 17, 21, 23)
ZooBank: http://zoobank.org/9B2C76D0-8E76-4A30-9E17-BE0EC12E6B44
Diagnosis. Distinguished among species of S. (Zygophleps) by the thumb-like process
of the subgenital plate (Fig. 13), and the small, robust, and unbranched right conjunc-
tival appendage with a carinated margin and surface with small spines on the second
half (Fig. 21).
Measurements: Male Holotype. Total length: 1.28 mm; pronotum length:
0.29 mm; pronotum width: 0.59 mm; scutellum length: 0.11 mm; abdominal width:
0.47 mm.
Material examined (n = 1): HOLOTYPE: BRAZIL: Pará: Ananindeua (adjoining
Marituba), 1.36583°S 48.37194°W, Nov 1963, Oliveira & Wygodzinsky, 1♂ (UCR_
ENT 00096701) (AMNH).
as in subgeneric description; sternum 6 strongly asymmetrical with a large triangular
posterolateral process with round apex strongly projecting from the anterolateral mar-
gin of subgenital plate (Fig. 13); subgenital plate weakly asymmetrical, posterior mar-
gin blunt (Fig. 13). GENITALIA: py as in subgeneric description; rp small, without a
shallow median groove, apex short, pointed and flattened dorsoventrally (Fig. 17); lp
as in subgeneric description; lca with the large projection about two times longer than
the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Distribution (Fig. 23). BRAZIL: Pará.
Etymology. Named for the dense setation of this species after the Latin “hirsutus”
meaning hairy.
Comments. Despite of the similar thumb-like process of the subgenital plate, which
resembles those in S. familia sp.n., S. orellanensis sp.n., and S. simla Emsley, this spe-
cies can be easily recognized by the distinct shape of the right paramere and right
conjunctival appendage.
26 DOI 10.1163/1876312X-bja10029
Schizoptera (Zygophleps) inaequalis sp.n.

(Figs 3, 6, 9, 13, 17, 21, 23)
ZooBank: http://zoobank.org/B8E8E2D7-2805-4580-93A9-30A9805CC912
Diagnosis. Distinguished among species of S. (Zygophleps) by the posteriorly pro-
jected and curved needle-shaped process of the subgenital plate (Fig. 13), and the long,
robust, and branched right conjunctival appendage with branches of different lengths
and width and armed with spines on apex (Fig. 21).
Material examined (n = 5): HOLOTYPE: BRAZIL: Pará: Belém: 8 km E
Belem, Ananindeua, 1.45297°S 48.56108°W, 06 Jun 1973, R.T. Schuh, 1♂ (UCR_
ENT 00096383) (AMNH). PARATYPES: BRAZIL: Pará: Belém: 8 km E Belem,
Ananindeua, 1.45297°S 48.56108°W, 02 Jun 1973, R.T. Schuh, 1♂ (UCR_ENT
00096405) (AMNH). ECUADOR: Orellana: Tiputini Biodiversity Sta. nr. Yasuni
National Park, 0.63194°S 76.14417°W, 220 m, 21 Oct 1998, T. L. Erwin et al., 1♂
(UCR_ENT 00028621) (USNM); 05 Feb 1999, 2♂ (UCR_ENT 00029119, UCR_
ENT 00029121) (USNM).
slightly lighter than remainder of hemelytra (Figure 3); rp with first half brown and
second half dark brown. SURFACE and VESTITURE: as in subgeneric description.
STRUCTURE: HEAD: rounded in lateral view (Fig. 9). THORAX: as in subgeneric
description. ABDOMEN: as in subgeneric description; sternum 6 asymmetrical with
a large triangular posterolateral process strongly projecting from the anterolateral mar-
gin of the subgenital plate (Fig. 13); subgenital plate weakly asymmetrical, posterior
margin blunt (Fig. 13). GENITALIA: py as in subgeneric description; rp large, with a
shallow median groove, flattened dorsoventrally, apex with two lateral projections (Fig.
17); lp as in subgeneric description; lca with one arm-like projection; v, ano, and at as
in subgeneric description.
Female. Unknown.
Distribution (Fig. 23). BRAZIL: Pará; ECUADOR: Orellana.
Etymology. Named for the unequal length of the branches of the right conjunctival
appendage after the Latin “inaequalis” meaning unequal.
Comments. See comments on S. aculeata sp.n.
Schizoptera (Zygophleps) inca sp.n.

(Figs 6, 9, 13, 18, 21, 23)
ZooBank: http://zoobank.org/3C7199DA-CABF-4AEE-9546-687E035680D9
Diagnosis. Distinguished among species of S. (Zygophleps) by the large, straight and
anterolaterally directed process of the subgenital plate with a very wide base (Fig. 13),
and the long, thin, and unbranched right conjunctival appendage without spines
(Fig. 21).
DOI 10.1163/1876312X-bja10029 27

Material examined (n = 2): HOLOTYPE: PERU: Madre de Dios: Los Amigos
Biol.Sta. trail 14, 12.57141°S 70.09538°W, 231 m, 22 Dec 2010, J. Heraty, 1♂,
(UCR_ENT 00086122) (UCR). PARATYPE: PERU: Madre de Dios: Same data as
holotype, 1♀ (UCR_ENT 00086123) (UCR).
Description. Male (Figs 6, 9). COLORATION: General coloration brown; C+Sc
slightly lighter than remainder of hemelytra; rp entirely pale brown. SURFACE and
VESTITURE: as in subgeneric description. STRUCTURE: HEAD: flat in lateral
description; sternum 6 weakly asymmetrical with a rounded posterolateral process
laterad-directed slightly projecting from the anterolateral margin of subgenital plate
(Fig. 13); subgenital plate strongly asymmetrical, posterior margin rounded (Fig. 13).
apex loong, slightly curved and pointed (Fig. 18); lp as in subgeneric description; lca
Female. As in subgeneric description (Fig. 11D–H).
Distribution (Fig. 23). PERU: Madre de Dios.
Etymology. Named for the Inca empire, which comprised a large territory that
arose from the Peruvian highlands and extended to Ecuador, Bolivia, Argentina, Chile,
and Colombia.
Comments. This is the only S. (Zygophleps) species where males and females are
known. Despite the large number of female specimens in our collection, we were una-
ble to match males and females based on morphology or locality, because females of
different species are likely very similar and several male-based species were collected at
the same locality. The male-female association of this species is based on molecular data
provided by Leon & Weirauch (2017). We were also able to document the egg of this
species (Fig. 11D, red ellipse).
Schizoptera (Zygophleps) kayi sp.n.

(Figs 3, 6, 9, 13, 18, 21, 23)
ZooBank: http://zoobank.org/54CF0AA7-B754-4D9B-A9E7-9DDF87699B53
Diagnosis. Distinguished among species of S. (Zygophleps) by the very short and
roughly quadrate process of the subgenital plate with apex slightly concave and with
anterior and posterior margins pointed (Fig. 13), and the small, thin, and unbranched
right conjunctival appendage with one small spine on the last third (Fig. 21).
76.14417°W, 220 m, 05 Feb 1999, T. L. Erwin et al., 1♂ (UCR_ENT 00028923)
(USNM).
28 DOI 10.1163/1876312X-bja10029

lateral view (Fig. 9). THORAX: as in subgeneric description. ABDOMEN: as in subge-
neric description; sternum 6 asymmetrical with a medium-sized triangular posterolat-
eral process strongly projecting from the anterolateral margin of subgenital plate (Fig.
13); subgenital plate weakly asymmetrical, posterior margin blunt. GENITALIA: py as
in subgeneric description; rp small, with a deep median groove, apex short, curved and
rounded (Fig. 18); lp as in subgeneric description; lca with the large projection about
Female. Unknown.
Etymology. Named in Honor of Andreas Kay who was a physicist, naturalist and
photographer and who dedicated his life to documenting the biodiversity of Ecuador.
Andreas Kay passed away in 2019 but made available more than 30.000 photographs
of Ecuadorian biodiversity.
Comments. See comments on S. erosa sp.n.
Schizoptera (Zygophleps) knyshovi sp.n.

(Figs 3, 6, 9, 13, 18, 21, 23)
ZooBank: http://zoobank.org/E2E33CEE-241C-49BA-92B8-66055A55625D
Diagnosis. Distinguished among species of S. (Zygophleps) by the “fishtail”-shaped
process of the subgenital plate (Fig. 13), and the long, robust, and unbranched right
conjunctival appendage without spines or with a small median spine on the margin
(Fig. 21).
Biodiversity Station near Yasuni National Park Erwin Transect - T/4, 0.63194°N
(USNM). PARATYPES: ECUADOR: Orellana: Same data as holotype, 4♂ (USNM)
(UCR_ENT 00028299, UCR_ENT 00028302, UCR_ENT 00028303, UCR_ENT
00028304) (USNM).
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: flat in lat-
eral view (Figure 9). THORAX: as in subgeneric description. ABDOMEN: as in
subgeneric description; sternum 6 weakly asymmetrical with a large triangular poste-
rolateral process strongly projecting from the anterolateral margin of subgenital plate
GENITALIA: py as in subgeneric description; rp small, with a deep median groove,
DOI 10.1163/1876312X-bja10029 29
description.
Female. Unknown.
Etymology. Named in Honor of Dr. Alexander Knyshov, a friend and expert
on Heteroptera, who has made important contributions to the knowledge of
Dipsocoromorpha.
Comments. This species shows variation of the right conjunctival appendage. Some
specimens have a spineless right conjunctival appendage while others have a small
median spine on the margin. Molecular data retrieved from specimens showing this
variation (Albuquerque et al., in preparation) support them as belonging to the same
species.
Schizoptera (Zygophleps) magnifica sp.n.

(Figs 3, 9, 13, 18, 21, 23)
ZooBank: http://zoobank.org/1CE349D0-7F66-4424-9F6E-CE3466ADEA4F
Diagnosis. Distinguished among species of S. (Zygophleps) by the large and roughly
quadrate process of the subgenital plate with anterior and posterior margins rounded
(Fig. 13), and the long, robust, and unbranched right conjunctival appendage with
medium-sized spines on the second half and apex enlarged (Fig. 21).
Material examined (n = 1): HOLOTYPE: COSTA RICA: Alajuela: Est. Biol.
Alberto Brenes, 10.21722°N 84.5975°W, 850 m, 03 Jul 1999, R.A. Anderson, 1♂
(UCR_ENT 00120217) (MTEC).
Description. Male (Figs 3, 9). COLORATION: General coloration brown; C+Sc
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: flat in lateral
description; sternum 6 asymmetrical with a small triangular posterolateral process
slightly projecting from the anterolateral margin of subgenital plate (Fig. 13); subgeni-
tal plate weakly asymmetrical, posterior margin rounded (Fig. 13). GENITALIA: py
as in subgeneric description; rp large, with a deep median groove, apex medium-sized,
strongly curved and rounded (Fig. 18); lp as in subgeneric description; lca with one
arm-like projection; v, ano, and at as in subgeneric description.
Female. Unknown.
Distribution (Fig. 23). COSTA RICA: Alajuela.
Etymology. Named for the beauty of this species after the Latin “magnificus” mean-
ing magnificent.
Comments. This species resembles S. solitaria sp.n. and S. ultima in having a similar
process of the subgenital plate. Nevertheless, these species can be distinguished by the
shape of the right conjunctival appendage.
30 DOI 10.1163/1876312X-bja10029
Schizoptera (Zygophleps) mallochi sp.n.

(Figs 3, 6, 9, 14, 18, 21, 23)
ZooBank: http://zoobank.org/DE0AE735-2192-44A6-8A64-AD6957358E59
Diagnosis. Distinguished among species of S. (Zygophleps) by the strongly asymmet-
rical subgenital plate with posterior margin blunt (Fig. 14), the “axe blade”-shaped
process of the subgenital plate with anterior margin pointed anterolaterally directed
and posterior margin rounded (Fig. 14), and the long, robust, and unbranched right
conjunctival appendage with a serrated margin (Fig. 21).
Biological Station, 10.43333°N 84.01667°W, 56 m, 30 Mar 1998, ALAS, Light Trap,
1♂ (UCR_ENT 00109206) (INBIO). PARATYPES: COSTA RICA: Heredia: Same
data as holotype, 1♂ (UCR_ENT 00109207) (INBIO); 16 Feb 1998, ALAS, Light
Trap, 1♂ (UCR_ENT 00109196) (INBIO); 16 Apr 1998, 2♂ (UCR_ENT 00109212,
UCR_ENT 00109215) (INBIO). Limon: Pococi, Colorado, Sector Cerro Cocoon 30
Km N Cariari, 10.7521°N 83.61356°W, 100 m, 10 Jan–11 Feb 1995, E. Rojas, 1♂
(UCR_ENT 00014513) (INBIO).
C+Sc concolorous with remainder of hemelytra (Fig. 3); rp almost entirely pale
brown with darker apex. SURFACE and VESTITURE: as in subgeneric descrip-
tion. STRUCTURE: HEAD: flat in lateral view (Fig. 9). THORAX: as in subgeneric
description. ABDOMEN: as in subgeneric description; sternum 6 asymmetrical with a
large triangular posterolateral process strongly projecting from the anterolateral margin
of subgenital plate (Fig. 14). GENITALIA: py as in subgeneric description; rp large,
with a shallow median groove, apex long, curved and rounded (Fig. 18); lp as in subge-
neric description; lca with the large projection about two times longer than the small;
v, ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named in Honor of Mr. John Malloch, who made important con-
tributions to the knowledge of Dipsocoromorpha and described the subgenus S.
(Zygophleps).
Comments. See comments on S. applanata sp.n.
Schizoptera (Zygophleps) mcateei sp.n.

(Figs 3, 6, 9, 14, 18, 21, 23)
ZooBank: http://zoobank.org/4A2E4068-D2A0-4C7F-9A59-A1CE5FA1C45B
Diagnosis. Distinguished among species of S. (Zygophleps) by the “fishtail”-shaped
process of the subgenital plate with anterior margin pointed and posterior margin
rounded (Fig. 14), and the large, robust, and unbranched right conjunctival append-
age with a serrated margin and surface with areolate sculptures (Fig. 21).
DOI 10.1163/1876312X-bja10029 31

Biological Station, 10.43333°N 84.01667°W, 56 m, 16 Apr 1998, ALAS, Light Trap,
1♂ (UCR_ENT 00109213) (INBIO).
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: rounded
subgeneric description; sternum 6 weakly asymmetrical with a large posterior process
projecting from the anterolateral margin of the subgenital plate (Fig. 14); subgenital
plate weakly asymmetrical, posterior margin rounded (Fig. 14). GENITALIA: py as
in subgeneric description; rp small, without a median groove, folded, apex pointed
(Fig. 18); lp as in subgeneric description; lca with the large projection about two times
Female. Unknown.
Etymology. Named in Honor of Dr. Waldo McAtee, who made important con-
tributions to the knowledge of Dipsocoromorpha and described the subgenus S.
(Zygophleps).
Comments. The right conjunctival appendage of Schizoptera (Zygophleps) mcateei
sp.n. is unique within the subgenus due to the presence of areolate sculptures on its
surface.
Schizoptera (Zygophleps) orellanensis sp.n.

(Figs 3, 6, 9, 14, 18, 21, 23)
ZooBank: http://zoobank.org/6DC73E87-A7C4-4D8F-A817-11AC2276F7D0
Diagnosis. Distinguished among species of S. (Zygophleps) by the small thumb-
like process of the subgenital plate (Fig. 14, black square), and the small, thin, and
unbranched right conjunctival appendage with a serrated margin (Figure 21).
Biodiversity Station nr Yasuni National Park, 0.63194°N 76.14417°W, 220 m, 26
Oct 1998, T. L. Erwin et al., 1♂ (UCR_ENT 00026824) (USNM). PARATYPES:
ECUADOR: Orellana: Tiputini Biodiversity Station Co.: nr Yasuni National Park
Erwin Transect - T/4, 0.63194°N 76.14417°W, 220 m, 24 Oct 1998, T. L. Erwin et
al., 2♂ (UCR_ENT 00028300, UCR_ENT 00028301) (USNM).
C+Sc lighter than remainder of hemelytra (Fig. 3); rp pale brown with darker apex.
as in subgeneric description; sternum 6 asymmetrical with a medium-sized triangu-
lar posterolateral process projecting from the anterolateral margin of subgenital plate
32 DOI 10.1163/1876312X-bja10029
apex medium-sized, weakly curved and rounded (Fig. 18); lp as in subgeneric descrip-
tion; lca with the large projection about three times longer than the small; v, ano, and
at as in subgeneric description.
Female. Unknown.
Etymology. Named for the locality where the species was collected.
Comments. See comments on S. hirsuta sp.n.
Schizoptera (Zygophleps) paraensis sp.n.

(Figs 3, 6, 9, 14, 18, 21, 23)
ZooBank: http://zoobank.org/27090C18-60B5-452D-95B2-A33FA77B1A17
Diagnosis. Distinguished among species of S. (Zygophleps) by the heart-shaped process
of the subgenital plate (Fig. 14), and the small, thin, and unbranched right conjuncti-
val appendage without spines (Fig. 21).
Material examined (n = 11): HOLOTYPE: BRAZIL: Pará: Paragominas,
Mineradora HYDRO [Norsk-Hydro bauxite mine] / 3.290222S, 47.718306W / 20
Nov–02 Dec 2018 / Col [= collectors]: Almeida, F. R. A. et al. / Armadilha lumi-
nosa CDC [= CDC light trap], 1♂ (UCR_ENT 00129753) (MPEG). PARATYPES:
BRAZIL: Pará: Same data as holotype, 5♂ (UCR_ENT 00129722, UCR_ENT
00129723, UCR_ENT 00129724, UCR_ENT 00129725, UCR_ENT 00129727)
(MPEG), 5♂ (UCR_ENT 00129728, UCR_ENT 00129729, UCR_ENT 00129730,
UCR_ENT 00129731, UCR_ENT 00129732) (UFPA).
lighter than remainder of hemelytra (Figure 3); rp entirely pale brown. SURFACE
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: flat in
lateral view (Fig. 9). THORAX: as in subgeneric description. ABDOMEN: as in
subgeneric description; sternum 6 weakly asymmetrical with a large triangular poste-
rolateral process strongly projecting from the anterolateral margin of subgenital plate
apex medium-sized, curved and pointed (Fig. 18); lp as in subgeneric description; lca
Female. Unknown.
Etymology. Named for the locality where the species was collected.
Comments. This species was collected in a secondary forest using CDC traps placed
on trees in open areas one meter above the ground. Specimens were collected on a min-
ing company area, in reclaimed forest, suggesting that this species may be adapted to
disturbed areas.
DOI 10.1163/1876312X-bja10029 33
Schizoptera (Zygophleps) peruviana sp.n.

(Figs 3, 6, 9, 14, 18, 21, 23)
ZooBank: http://zoobank.org/37CF8B04-7EE3-41CB-A84E-3C4DF0C756FD
process of the subgenital plate with anterior and posterior margins rounded (Fig. 14),
and the long, robust, and unbranched right conjunctival appendage with a long spine
on the margin and surface composed of small and medium-sized spines on the second
half (Fig. 21).
Material examined (n = 1): HOLOTYPE: PERU: Loreto: Rio Sucusari Co.: 160km
NE Iquitos, Explornapo Camp, Rio Sucusari 2 km from Rio Napo, 3.74367°S
73.25163°W, 27–31 Aug 1992, P. E. Skelley, 1♂ (UCR_ENT 00012012) (FSCA).
lighter than remainder of hemelytra (Figure 3); rp entirely pale brown. SURFACE
generic description; sternum 6 weakly asymmetrical with a small triangular postero-
lateral process projecting from the anterolateral margin of subgenital plate (Fig. 14);
subgenital plate weakly asymmetrical, posterior margin blunt (Fig. 14). GENITALIA:
py as in subgeneric description; rp small, with a deep median groove, apex long, not
tapering, weakly curved and rounded (Fig. 18); lp as in subgeneric description; lca
Female. Unknown.
Distribution (Fig. 23). PERU: Loreto.
Etymology. Named for the country where the species was collected.
Comments. See comments on S. erwini sp.n.
Schizoptera (Zygophleps) pichincha sp.n.

(Figs 3, 6, 9, 14, 18, 21, 23)
ZooBank: http://zoobank.org/6249FA40-2555-4C4D-B4FE-0B12101437A1
Diagnosis. Distinguished among species of S. (Zygophleps) by the process of the sub-
genital plate with anterior margin well-developed and rounded and posterior margin
with a distinct spur (Fig. 14), and the long, robust, and unbranched right conjunctival
appendage with a serrated margin (Fig. 21).
Material examined (n = 2): HOLOTYPE: ECUADOR: Pichincha: Reserva
Palenque, 0.27696S, 78.29946W, 01 Jul 1989–15 Jul 1989, malaise trap, 1♂ (UCR_
ENT 00091355) (FSCA). PARATYPE: ECUADOR: Pichincha: Same data as holo-
type, 1♂ (UCR_ENT 00091336) (FSCA).
34 DOI 10.1163/1876312X-bja10029

C+Sc lighter than remainder of hemelytra (Fig. 3); rp brown with apex dark brown.
as in subgeneric description; sternum 6 slightly asymmetrical with a small triangular
posterolateral process slightly projecting from the anterolateral margin of subgenital
14). GENITALIA: py as in subgeneric description; rp large, without a median groove,
folded, apex pointed and laterally directed in dorsal view [not shown in the drawing]
(Fig. 18); lp as in subgeneric description; lca with the large projection about two times
Female. Unknown.
Distribution (Fig. 23). ECUADOR: Pichincha.
Etymology. Named after the locality where the species was collected.
Comments. See comments on S. bianca sp.n.
Schizoptera (Zygophleps) pinduca sp.n.

(Figs 3, 6, 9, 14, 19, 22, 23)
ZooBank: http://zoobank.org/11729F32-D337-4281-8B90-D0BC3621FF63
Diagnosis. Distinguished among species of S. (Zygophleps) by the spatulated process of
the subgenital plate with anterior margin rounded and posterior margin pointed, lat-
erally well-projected (Fig. 14), and the long, thin, and unbranched right conjunctival
appendage with a serrated margin and surface with short serrate lines (Fig. 22).
Material examined (n = 1): HOLOTYPE: BRAZIL: Pará: Paragominas,
Mineradora HYDRO [Norsk-Hydro bauxite mine] / 3.290222S, 47.718306W / 20
Nov–02 Dec 2018 / Col [= collectors]: Almeida, F. R. A. et al. / Armadilha luminosa
CDC (20m) [= CDC light trap], 1♂ (UCR_ENT 00129726) (MPEG).
Description. Male. Body length: 1.20 mm (Figs 3, 6, 9); COLORATION: General
coloration brown; C+Sc lighter than remainder of hemelytra (Fig. 3); rp entirely pale
HEAD: flat in lateral view (Fig. 9). THORAX: as in subgeneric description.
ABDOMEN: as in subgeneric description; sternum 6 asymmetrical with a large pos-
terolateral process strongly projecting from the anterolateral margin of the subgenital
plate (Fig. 14); subgenital plate asymmetrical, posterior margin rounded (Fig. 14).
Female. Unknown.
Etymology. Named as an homage to Aurino Quirino Gonçalves, best known as
Pinduca, who represents the local culture through his music. He is a musician and
DOI 10.1163/1876312X-bja10029 35
composer from the north of Brazil famous for Carimbó, a traditional dance from the
state of Pará, Brazil, which has African, indigenous, and European roots, and is one of
the cultural icons of this region.
Comments. This species was collected in a secondary forest using CDC traps
installed 20 meters above the ground, suggesting that these insects are capable of flight
at relatively high heights despite their minute size.
Schizoptera (Zygophleps) prima sp.n.

(Figs 3, 6, 9, 14, 19, 22, 23)
ZooBank: http://zoobank.org/55AA198E-63EE-4ED8-BD74-949F19741433
Diagnosis. Distinguished among species of S. (Zygophleps) by the trapezoidal process
of the subgenital plate with anterior margin longer than the posterior margin (Fig.
14), and the long, thin, and unbranched right conjunctival appendage with a serrated
margin (Fig. 22).
76.14417°W, 220 m, 05 Feb 1999, T. L. Erwin et al., 1♂ (UCR_ENT 00027927)
(USNM). PARATYPES: ECUADOR: Orellana: Same data as holotype, 2♂ (UCR_
ENT 00027928, UCR_ENT 00028983) (USNM).
Description. Male (Figes 3, 6, 9). COLORATION: General coloration pale brown;
generic description; sternum 6 weakly asymmetrical with a large pointed posterolateral
process strongly projecting posteriorly (Fig. 14); subgenital plate weakly asymmetrical,
posterior margin rounded (Fig. 14). GENITALIA: py as in subgeneric description; rp
large, with a deep median groove, apex long, thin, slightly curved and pointed (Fig.
19); lp as in subgeneric description; lca with the large projection about two times
Female. Unknown.
Etymology. Named for the fact that this was the first species of S. (Zygophleps) that
the first author sorted after the Latin “primus” meaning first.
Comments. The process of the subgenital plate is a distinctive character among
other species of this subgenus.
Schizoptera (Zygophleps) priva sp.n.

(Figs 3, 6, 9, 14, 19, 22, 23)
ZooBank: http://zoobank.org/8FF5D525-62CD-4210-9498-4610BC668E6C
Diagnosis. Distinguished among species of S. (Zygophleps) by the rectangular pro-
cess of the subgenital plate with a median concavity (Fig. 14, black arrow) and with
36 DOI 10.1163/1876312X-bja10029
anterior and posterior margins rounded (Fig. 14), and the short, thin, and unbranched
right conjunctival appendage with a serrated margin (Fig. 22).
Biological Station, 10.43333°N 84.01667°W, 56 m, 05 Jan 2000, ALAS, Eugenia sp.
(Myrtaceae), 1♂ (UCR_ENT 00109296) (INBIO).
and VESTITURE: as in subgeneric description. STRUCTURE: HEAD: rounded
subgeneric description; sternum 6 weakly asymmetrical with a small triangular pos-
terolateral process projecting from the anterolateral margin of the subgenital plate
large projection about three times longer than the small; v, ano, and at as in subgeneric
description.
Female. Unknown.
Etymology. Named for the fact that this species is known just from the holotype
after the Latin “privus” meaning single.
Comments. See comments on S. erwini sp.n.
Schizoptera (Zygophleps) ramosa sp.n.

(Figs 4, 7, 10, 14, 19, 22, 23)
ZooBank: http://zoobank.org/C4CBDC22-6E04-422E-A1AF-38563D5C6402
Diagnosis. Distinguished among species of S. (Zygophleps) by the claw-shaped process
of the subgenital plate posteriorly directed (Fig. 14), the right paramere large, with-
out a median groove and with a pointed apex forming a 90º angle to the rest of the
stem (Fig. 19), and the small, thin, and branched right conjunctival appendage with
branches of same length and armed with spines on apex (Fig. 22).
Material examined (n = 2): HOLOTYPE: ECUADOR: Orellana: Northern
Production Facility, Reserva Etnica Waorani, 0.65714°N 76.453°W, 216 m, 23
Feb 1995, T. L. Erwin et al., 1♂ (UCR_ENT 00028048) (USNM). PARATYPE:
BRAZIL: Pará: Ananindeua (adjoining Marituba), 1.36583°S 48.37194°W, Nov
1963, Oliveira & Wygodzinsky, 1♂ (UCR_ENT 00096649) (AMNH).
C+Sc slightly lighter than remainder of hemelytra (Fig. 4); basal half of rp pale brown,
distal half dark brown. SURFACE and VESTITURE: as in subgeneric description.
STRUCTURE: HEAD: flat in lateral view (Fig. 10). THORAX: as in subgeneric
DOI 10.1163/1876312X-bja10029 37
description. ABDOMEN: as in subgeneric description; sternum 6 asymmetrical with

a medium-sized triangular posterolateral process projecting from the anterolateral mar-
gin of subgenital plate (Fig. 14); subgenital plate weakly asymmetrical, posterior mar-
gin blunt (Fig. 14). GENITALIA: py as in subgeneric description; lp as in subgeneric
Female. Unknown.
Distribution (Fig. 23). BRAZIL: Pará; ECUADOR: Orellana.
Etymology. Named for the branched right conjunctival appendage after the Latin
“ramosus” meaning branched.
Comments. See comments on S. corallia Emsley.
Schizoptera (Zygophleps) simla Emsley, 1969

Schizoptera (Zygophleps) simla Emsley, 1969: 25, 66 (orig. descr.)
(Figs 4, 7, 10, 14, 19, 22, 23)
Updated Diagnosis. Distinguished among species of S. (Zygophleps) by the thumb-
like process of the subgenital plate (Fig. 14), and the long, robust, and unbranched
right conjunctival appendage with spines on apex (Fig. 22).
Measurements: Paratype. Total length: 1.39 mm; pronotum length: 0.26 mm; pro-
notum width: 0.58 mm; scutellum length: 0.13 mm; abdominal width: 0.48 mm.
Valley: William Beebe Tropical Research Station, at light, 28 Sep 1961, M. G.
Emsley, 1♂ (AMNH_IZC 00150720) (AMNH). PARATYPES: TRINIDAD AND
TOBAGO: Arima Valley: Same data as holotype except 12 Oct 1961 and 12 Jan1962,
4♂ (AMNH_IZC 00343395, AMNH_IZC 00343396) (AMNH) (AMNH, USNM,
two slides, two glycerine). ADDITIONAL MATERIAL: SURINAME: Brokopondo:
Brownsberg Nature Park Camp, 4.948733N, 55.180900W, 25 Aug–02 Sep 2007, G.
Kung, A. Kreuter (UCR_ENT 00086068) (UCR). BRAZIL: Pará: Paragominas: M
ineradora HYDRO [Norsk-Hydro bauxite mine], 3.290222S, 47.718306W, 16 Sep
2018, Col Almeida, F. R. A. et al., Armadilha luminosa CDC (10m), 1♂ (UCR_ENT
00129733) (MPEG); FRENCH GUIANA: Roura: Amazon Nature Lodge, 30 Km SE
Roura on Kaw Rd., 4.559500N, 52.205567W, 2–8 Jun 2005, UV trap, J. E. Eger &
M. T. Messenger (UCR_ENT 00012001) (FSCA)
Re-description. Male (Figs 4, 7, 10). COLORATION: General coloration
brown; C+Sc slightly lighter than remainder of hemelytra (Fig. 4); rp entirely pale
medium-sized rounded posterolateral process projecting from the anterolateral margin
blunt (Fig. 14). GENITALIA: py as in subgeneric description; rp large, with a deep
median groove, apex short, strongly curved and pointed (Fig. 19); lp as in subgeneric
38 DOI 10.1163/1876312X-bja10029
Female. Unknown.
Distribution (Fig. 23). BRAZIL: Pará; FRENCH GUIANA: Roura; SURINAME:
Brokopondo; TRINIDAD AND TOBAGO: Arima Valley.
Comments. Emsley (1969) described this species from Trinidad based on male
specimens. We here redescribe S. simla Emsley and expand its distribution to Brazil,
French Guiana, and Suriname. The Brazilian specimen was collected in a secondary
forest using CDC traps installed ten meters above the ground. This species resembles
S. familia sp.n. in having a similar process of the subgenital plate and right conjunc-
tival appendage. However, they can be distinguished by the presence or absence of a
median spine on the margin of the right conjunctival appendage. The spine is absent
in S. simla, while it is present in S. familia sp.n. Additionally, S. familia sp.n. is only
recorded from Ecuador.
Schizoptera (Zygophleps) solitaria sp.n.

(Figs 4, 7, 14, 19, 22, 23)
ZooBank: http://zoobank.org/71569DD4-60DD-4439-9EFE-D27720723069
Diagnosis. Distinguished among species of S. (Zygophleps) by the roughly quadrate
and very large process of the subgenital plate with anterior and posterior margins
rounded (Fig. 14), and the long, thin, and unbranched right conjunctival appendage
with a serrated margin and spines on apex (Fig. 22).
Material examined (n = 1): HOLOTYPE: BOLIVIA: La Paz: La Paz, Alto Beni,
16.5°S 68.15°W, 1600 m, 12–15 Apr 2003, B. Brown, S. Thompson, and E. Zumbado,
1♂ (UCR_ENT 00086525) (LACM).
Description. Male. Body length: 1.27 to 1.52 mm (Figs 4, 7); COLORATION:
General coloration brown; C+Sc slightly lighter than remainder of hemelytra (Fig.
4); rp entirely pale brown. SURFACE and VESTITURE: as in subgeneric descrip-
tion. STRUCTURE: THORAX: as in subgeneric description. ABDOMEN: as in
subgeneric description; sternum 6 weakly asymmetrical with a small posterolateral
process projecting from the anterolateral margin of sternum 7 (Fig. 14); subgenital
plate asymmetrical, posterior margin rounded (Fig. 14). GENITALIA: py as in subge-
neric description; rp small, with a deep median groove, apex long, slightly curved and
rounded (Fig. 19); lp as in subgeneric description; lca with the large projection about
Female. Unknown.
Distribution (Fig. 23). BOLIVIA: La Paz.
Etymology. Named for the fact that this species is known just from the holotype
after the Latin “solitarius” meaning alone.
Comments. See comments on S. magnifica sp.n.
DOI 10.1163/1876312X-bja10029 39
Schizoptera (Zygophleps) speira sp.n.

(Figs 4, 7, 10, 15, 19, 22, 23)
ZooBank: http://zoobank.org/80DF0B7B-20A4-4DCC-960B-4E1C6045DD1A
Diagnosis. Distinguished among species of S. (Zygophleps) by the thin and elongated
process of the subgenital plate with anterior margin projecting anterolaterally (Fig.
15), and the extremely long, robust, and coiled right conjunctival appendage forming
about 1 turn, with large spines on the surface (Fig. 22).
width: 0.61 mm; scutellum length: 0.12mm; abdominal width: 0.48 mm.
Material examined (n = 1): HOLOTYPE: ECUADOR: Orellana: Waorani, 1km
S. Onkone Gare Camp, 0.63333°N 76.6°W, 220 m, 25 Jun 1994, T. L. Erwin et al.,
1♂ (UCR_ENT 00026409) (USNM).
as in subgeneric description; sternum 6 asymmetrical with a large triangular poste-
rolateral process projecting from the anterolateral margin of the subgenital plate
Female. Unknown.
Etymology. Named for the shape of the right conjunctival appendage after the
Greek “speira” meaning spiral.
Comments. This species can be easily recognized among species of S. (Zygophleps)
by the unique shape of the right conjunctival appendage. Additionally, see comments
on S. emsleyi sp.n.
Schizoptera (Zygophleps) stricta sp.n.

(Figs 4, 7, 10, 15, 19, 22, 23)
ZooBank: http://zoobank.org/8479D43D-1E71-4D99-AC38-79EEF89CE945
Diagnosis. Distinguished among species of S. (Zygophleps) by the long, straight, and
apically rounded process of the subgenital plate (Fig. 15), and the large, robust, and
branched right conjunctival appendage on apex with branches of different lengths and
without spines (Fig. 22).
Measurements: Male Holotype. Total length: 1.33 mm; pronotum length:
0.20 mm; pronotum width: 0.53 mm; scutellum length: 0.12 mm; abdominal width:
0.48 mm.
40 DOI 10.1163/1876312X-bja10029
Material examined (n = 6): HOLOTYPE: ECUADOR: Orellana: Transect

Ent. 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, Onkone Gare Camp,
0.65714°N 76.453°W, 216 m, 03 Oct 1996, T. L. Erwin et al., 1♂ (UCR_ENT
00028088) (USNM). PARATYPES: ECUADOR: Orellana: Res. Ethnica Waorani,
1km S. Onkone Gare Camp, 0.63333°N 76.6°E, 220 m, 23 Jan 1994, T. L. Erwin et
al., 1♂ (UCR_ENT 00026383) (USNM); 25 Jan 1994, 1♂ (UCR_ENT 00026380)
(USNM); 21 Jun 1994, 1♂ (UCR_ENT 00026396) (USNM); Tiputini Biodiversity
Station nr Yasuni National Park Erwin Transect - T/9, 0.63194°N 76.14417°W,
220 m, 05 Feb 1999, T. L. Erwin et al., 2♂ (UCR_ENT 00028812, UCR_ENT
00028982) (USNM).
C+Sc concolorous with remainder of hemelytra (Fig. 4); rp pale brown with apex dark
small rounded posterolateral process projecting from the anterolateral margin of sub-
genital plate (Fig. 15). GENITALIA: py as in subgeneric description; rp large, with a
deep median groove, apex long, slightly curved, and remarkably rounded (Fig. 19); lp
as in subgeneric description; lca with the large projection about three times longer than
the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Etymology. Named for the straight process of the subgenital plate after the Latin
“strictus” meaning straight.
Comments. The process of the subgenital plate and the right conjunctival append-
age in this species are easily recognized among species of S. (Zygophleps).
Schizoptera (Zygophleps) ultima Emsley, 1969

Schizoptera (Zygophleps) ultima Emsley, 1969: 25, 67 (orig. descr.)
(Figs 15, 19, 22, 23)
Updated diagnosis. Distinguished among species of S. (Zygophleps) by the roughly
quadrate and very large process of the subgenital plate with anterior margin rounded
and posterior margin pointed (Fig. 15), and the long, thin, and unbranched right con-
junctival appendage with a serrated margin, almost forming a whole turn (Figure 22).
Valley: William Beebe Tropical Research Station, at light, 02 Nov 1961, M. G. Emsley,
1♂ (AMNH_IZC 00150336) (AMNH) (slide mounted). PARATYPE: TRINIDAD
AND TOBAGO: Arima Valley: Same data as holotype except for 29 Dec 1961, 1♂
(USNM).
Re-description. Male. Body length: 1.50 mm; COLORATION: General coloration
brown; C+Sc concolorous with remainder of hemelytra. SURFACE and VESTITURE:
as in subgeneric description. STRUCTURE: THORAX: as in subgeneric description.
DOI 10.1163/1876312X-bja10029 41

medium-sized posterolateral rounded process projecting from the anterolateral mar-
gin of subgenital plate (Fig. 15); subgenital plate asymmetrical, posterior margin
rounded (Fig. 15). GENITALIA: py as in subgeneric description; rp with a strongly
deep median groove, apex medium-sized, slightly curved and pointed [missing in the
holotype] (Fig. 19); lp as in subgeneric description; lca with the large projection about
three times longer than the small; v, ano, and at as in subgeneric description.
Female. Unknown.
Distribution (Fig. 23). TRINIDAD AND TOBAGO: Arima Valley.
Comments. Emsley (1969) described this species from Trinidad based on two male
specimens. He mentioned the remarkably long and serrated right conjunctival append-
age that sets this species apart from other species of S. (Zygophleps). Additionally, this
species has a peculiar process of the subgenital plate, which is roughly quadrate and
remarkably enlarged, slightly similar to those of S. magnifica sp.n. and S. solitaria sp.n.
Schizoptera (Zygophleps) unica McAtee & Malloch, 1925

Schizoptera (Zygophleps) unica McAtee & Malloch, 1925: 67, 18 (orig. descr.)
(Figs 4, 7, 10, 15, 23)
Updated Diagnosis. Distinguished among species of S. (Zygophleps) by the weakly
asymmetrical sternum 6 with a small rounded posterior process strongly projecting
from the anterior margin of the subgenital plate (Fig. 15), the weakly asymmetrical
subgenital plate (Fig. 15), the thumb-like process of the subgenital plate (Fig. 15).
Material examined (n = 1): HOLOTYPE: GUATEMALA: Livingston: 07 May
19??, Barber & Schwarz (UCR_ENT 00028619) (USNM).
Re-description. Male (Figs 4, 7, 10). COLORATION: General coloration dark
brown; C+Sc concolorous with remainder of hemelytra (Fig. 4). SURFACE and
VESTITURE: as in subgeneric description. STRUCTURE: HEAD: rounded in lateral
description; subgenital plate weakly asymmetrical, posterior margin rounded (Fig. 15);
process of the subgenital plate thumb-like (Fig. 15). GENITALIA: unknown.
Female. Unknown.
Distribution (Fig. 23). GUATEMALA: Livingston.
Comments. McAtee & Malloch (1925) described this species based on a single
male collected in Guatemala and did not document genitalic features. The holotype
is point-mounted, and we were unable to access genitalic features. For this reason,
the redescription is uniquely based on somatic features. Moreover, Emsley (1969)
expanded the original description, illustrated the subgenital plate and male genitalia of
S. unica based on specimens he collected in Trinidad. However, after careful examina-
tion of Emsley’s specimens and comparison with the original drawing and the holotype
of S. unica, we concluded that the specimens which Emsley collected and documented
do not belong to S. unica.
42 DOI 10.1163/1876312X-bja10029
Fig. 2. Habitus images of S. (Zygophleps) spp. in dorsal view. Scale bar: 0.5 mm.
DOI 10.1163/1876312X-bja10029 43
44 DOI 10.1163/1876312X-bja10029
DOI 10.1163/1876312X-bja10029 45
Fig. 5. Habitus images of S. (Zygophleps) spp. in ventral view. Scale bar: 0.5 mm.
46 DOI 10.1163/1876312X-bja10029
DOI 10.1163/1876312X-bja10029 47
48 DOI 10.1163/1876312X-bja10029
Fig. 8. Habitus images of S. (Zygophleps) spp. in left lateral view. Scale bar: 0.5 mm.
DOI 10.1163/1876312X-bja10029 49
50 DOI 10.1163/1876312X-bja10029
DOI 10.1163/1876312X-bja10029 51
Fig. 11. Female of S. (Zygophleps) inca sp.n. A–C Habitus images. A. Dorsal view. B. Ventral view. C.
Lateral view. D–G Abdomen. D. Ventral view (red arrow showing the silhouette of an egg; blue arrow
showing the silhouette of the spermatheca). E. Ventral view. F. Dorsal view. G. Right lateral view. H.
Spermatheca. Abbreviations: at = anal tube, spd = spermathecal duct, spgl = spermathecal gland, spgld =
spermathecal gland duct, spr = spermathecal reservoir, st7 = sternite 7, t7 = tergite 7, t8 = tergite 8, t9 =
tergite 9. Scale bars: 0.5 mm (A–C); 0.1 mm (D–H).
52 DOI 10.1163/1876312X-bja10029
Fig. 12. Images of abdominal apex showing the diagnostic features of sternum 6 and the subgenital plate
(sternum 7) of S. (Zygophleps) spp., ventral view (black arrow showing the tooth-like projection; black
rectangle showing the morphological variation of the process of the subgenital plate). Scale bar: 0.1 mm.
DOI 10.1163/1876312X-bja10029 53
(sternum 7) of S. (Zygophleps) spp., ventral view. Scale bar: 0.1 mm.
54 DOI 10.1163/1876312X-bja10029
Fig. 14. Images of abdominal apex showing the diagnostic features of sternum 6 and the subgenital
plate (sternum 7) of S. (Zygophleps) spp., ventral view (black square showing the thumb-like process of the
subgenital plate; black arrow showing the median concavity). Scale bar: 0.1 mm.
DOI 10.1163/1876312X-bja10029 55
(sternum 7) of S. (Zygophleps) spp., ventral view. Scale bar: 0.1 mm.
56 DOI 10.1163/1876312X-bja10029
Fig. 16. Line drawings of the right paramere of S. (Zygophleps) spp. Scale bars: 0.05 mm.
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Fig. 20. Line drawings of the right conjunctival appendage of S. (Zygophleps) spp. Scale bars: 0.05 mm.
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Fig. 23. Distribution records of S. (Zygophleps) spp.
Conclusions
Molecular phylogenetic analyses of Schizoptera show that S. (Zygophleps) is mono-
phyletic (Leon & Weirauch 2017; Albuquerque et al., in prep.). Despite the ten-fold
increase of described species in this subgenus, the triangular shape of the posterior
membranal cell (Fig. 1A) remains an excellent diagnostic feature for this taxon. Species
of this subgenus are characterized by very similar habitus and somatic morphological
features, which can make identification difficult. However, the morphological diversity
of the male subgenital plate and genitalia are astounding in this subgenus and provide
reliable species-diagnostic characters. In particular, the shape of the process of the sub-
genital plate, right paramere and right conjunctival appendage are crucial in determin-
ing species-level identification.
Our taxonomic revision of S. (Zygophleps) considerably improved our knowledge of
the distribution patterns of this group. Despite the substantial number of new species
for this taxon, several of these species were collected in the same locality (particularly
Heredia and Orellana in Costa Rica and Ecuador, respectively) (Fig. 23). The patterns
64 DOI 10.1163/1876312X-bja10029
herein observed along with field observations and collection data suggest that S.
(Zygophleps) species might be active fliers. Ultimately, the distributional gaps observed
are likely due to the lack of material from many areas in Central and South America.
Acknowledgments
We thank Dr. Alexander Knyshov (UCR) and Rochelle Hoey-Chamberlain (UCR) for
their extremely helpful comments during the development of this project. Thanks to
Caroline Souza and an anonymous reviewer for their valuable comments on the manu-
script. The authors are grateful the Brazilian Conselho Nacional de Desenvolvimento
Científico e Tecnológico-CNPq for financial support (CNPq UNIVERSAL
420294/2016-3, FRAA 140079/2018-1, JAMF 311345/2018-2) and The Comissão
Fulbright Brasil. We also thank the Norsk-Hydro bauxite mine and the Biodiversity
Research Consortium Brazil-Norway (BRC) for support. This project is part of the
“ARTS: Litter Bugs: revisionary and phylogenetic research on the least studied true
bug infraorder (Insecta: Hemiptera: Dipsocoromorpha)” project supported by the
National Science Foundation DEB-1257702.
References
Almeida, F.R.A., Carvalho-Filho, F.S. & Fernandes, J.A.M. (2019) A new species of Hypsohapsis
(Heteroptera: Schizopteridae: Hypselosomatinae) from the Brazilian Amazon. Zootaxa 4551 (4):
463–470.
Emsley, M.G. (1969) The Schizopteridae (Hemiptera: Heteroptera) with the descriptions of new species
from Trinidad. Memoirs of the American Entomological Society 25: 1–154.
Knyshov, A., Hoey-Chamberlain, R. & Weirauch, C. (2018) Comparative morphology of male genitalic
structures in the minute litter bugs Dipsocoromorpha (Insecta: Hemiptera: Heteroptera). Journal of
Morphology 279: 1480–1517.
Knyshov, A., Weirauch, C. & Hoey-Chamberlain, R. (2020) Phylogenetic relationships and revised clas-
sification of the true bug infraorder Dipsocoromorpha (Insecta: Hemiptera: Heteroptera). Cladistics
1–28.
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(Odontorhagus) reveals vast undocumented biodiversity in the largest litter bug genus Schizoptera
Fieber (Hemiptera: Dipsocoromorpha). Zootaxa 4184 (2): 255–284.
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Weirauch, C. & Fernandes, J.A.M. (2015) The minute litter bugs (Dipsocoromorpha). In: Panizzi,
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(Hemiptera: Schizopteridae). Zootaxa 4370 (2): 156–170.

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Insect Systematics & Evolution (2022) DOI 10.

Taxonomic revision of the minute litter bug subgenus

Flavio Albuquerquea,b,*,**, Jose A.M. Fernandesa,*** and Christiane Weirauchb,****

© Koninklijke Brill NV, Leiden, 2022DOI 10.1163/1876312X-bja10029

Material and nethods

Images and line drawings

compound microscope (Oberkochen, Germany) using Archimed V5.4.1 (Microvision).

Key to the species of Schizoptera (Zygophleps) based on males

2. Right conjunctival appendage branched (Figs 20, 22 e.g., S. aculeata sp.n., S.

Schizoptera (Zygophleps) aculeata sp.n.

Schizoptera (Zygophleps) applanata sp.n.

GENITALIA: py as in subgeneric description; rp large, with a deep median groove,

Schizoptera (Zygophleps) bella sp.n.

Schizoptera (Zygophleps) biancae sp.n.

Schizoptera (Zygophleps) cassiae sp.n.

Schizoptera (Zygophleps) confusa sp.n.

Measurements: Holotype. Total length: 1.19 mm; pronotum length: 0.24 mm;

Schizoptera (Zygophleps) corallia Emsley, 1969

Schizoptera (Zygophleps) costarriquenha sp.n.

Description. Male (Figs 2, 5, 8). COLORATION: General coloration brown; C+Sc

Schizoptera (Zygophleps) cuspida sp.n.

posterolateral process slightly projecting from the anterolateral margin of subgenital

Schizoptera (Zygophleps) ecuadoriana sp.n.

Schizoptera (Zygophleps) elegantissima sp.n.

Schizoptera (Zygophleps) emsleyi sp.n.

Measurements: Total length: 1.36 mm; pronotum length: 0.18 mm; pronotum

Schizoptera (Zygophleps) erosa sp.n.

SURFACE and VESTITURE: as in subgeneric description. STRUCTURE: HEAD:

Schizoptera (Zygophleps) erwini sp.n.

Schizoptera (Zygophleps) falcata sp.n.

Schizoptera (Zygophleps) familia sp.n.

Schizoptera (Zygophleps) furcata sp.n.

Measurements: Total length: 1.49 mm; pronotum length: 0.20 mm; pronotum

Schizoptera (Zygophleps) fuscodorsata sp.n.

GENITALIA: py as in subgeneric description; rp small, with a shallow median groove,

Schizoptera (Zygophleps) hirsuta sp.n.

Schizoptera (Zygophleps) inaequalis sp.n.

Schizoptera (Zygophleps) inca sp.n.

Measurements: Holotype. Total length: 1.26 mm; pronotum length: 0.19 mm;

Schizoptera (Zygophleps) kayi sp.n.

Description. Male (Figs 3, 6, 9). COLORATION: General coloration brown; C+Sc

Schizoptera (Zygophleps) knyshovi sp.n.

Schizoptera (Zygophleps) magnifica sp.n.

Schizoptera (Zygophleps) mallochi sp.n.

Schizoptera (Zygophleps) mcateei sp.n.

Measurements: Total length: 1.20 mm; pronotum length: 0.19 mm; pronotum

Schizoptera (Zygophleps) orellanensis sp.n.

Schizoptera (Zygophleps) paraensis sp.n.

Schizoptera (Zygophleps) peruviana sp.n.

Schizoptera (Zygophleps) pichincha sp.n.

Description. Male (Figs 3, 6, 9). COLORATION: General coloration brown;

Schizoptera (Zygophleps) pinduca sp.n.

Schizoptera (Zygophleps) prima sp.n.

Schizoptera (Zygophleps) priva sp.n.

Schizoptera (Zygophleps) ramosa sp.n.

description. ABDOMEN: as in subgeneric description; sternum 6 asymmetrical with

Schizoptera (Zygophleps) simla Emsley, 1969

Schizoptera (Zygophleps) solitaria sp.n.

Flavio Albuquerquea,b,,, Jose A.M. Fernandesa, and Christiane Weirauchb,****

© Koninklijke Brill NV, Leiden, 2022DOI 10.1163/1876312X-bja10029